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BULLETIN OF

THE BRITISH MUSEUM
(NATURAL HISTORY)

ZOOLOGY

Vol. 21

19701972

BRITISH MUSEUM (NATURAL HISTORY)
LONDON: 1975

DATES OF PUBLICATION OF THE PARTS

No. I . . . -.22 December 1970

No. 2 . . . . .28 April 1971

No. 3 . . . . .10 September 1971

No. 4 . . . .10 September 1971

No. 5 . . . .21 September 1971

No. 6 . . . . .3 December 1971

No. 7 . . . .3 December 1971

No. 8 ..... 4 February 1972

Printed in Great Britain by John Wright and Sons Ltd. at The Stonebridge Press, Bristol BS4 5NU

CONTENTS

ZOOLOGY VOLUME 21

No. i. Decapod crustacean larvae collected during the International
Indian Ocean Expedition. Families Raninidae and Homolidae.
By A. L. RICE

No. 2. Bats from Ethiopia collected by the Great Abbai Expedition,
1968. By J. E. HILL and P. MORRIS (Pis. 1-3)

No. 3. On some species of parasitic worms in the 'Discovery' collections
obtained in the years 1925-1936. By S. MARKOWSKI

No. 4. Mites of the genus Hypoaspis Canestrini, 1884 5. sir. and related
forms (Acari : Mesostigmata) associated with beetles. By
M. COSTA ..........

No. 5. The polychaete fauna of the Solomon Islands. By P. E. GIBBS

No. 6. Lizards and snakes from Transjordan, recently acquired by the
British Museum (Natural History). By Y. L. WERNER (Pis. 1-6)

No. 7. Conchoecia from the North Atlantic, the 'procera' group. By
M. V. ANGEL

Miscellanea ..........

Conchoecia pseudoparthenoda (nov. sp.), a new halocyprid ostra-

cod for the Tropical North Atlantic. By M. V. ANGEL .

The type specimens and identity of the species described in the

genus Lithobius by George Newport in 1844, 1845 and 1849

(Chilopoda, Lithobiomorpha). By E. H. EASON .

A re-examination of the chestnut-shouldered wren complex of

Australia. By C. J. O. HARRISON .....

A new species of Lironeca (Isopoda ;

on cichlid fishes in Lake Tanganyika.

Eunice manihine sp. nov. (Polychaeta

of the flavus-bidentate group from

Indian Ocean. By M. R. LONGBOTTOM ....

Original description of the gangetic dolphin, Platanista gange-
tica, attributed to William Roxburgh. By G. PILLERI .
A new genus and species of Sudan leech formerly confused with
Limnatis nilotica (Hirudinidae s.l. : Hirudinea). By L. R.
RICHARDSON .........

The type-species of the genera Phoxinellus, Pseudophoxinus and
Paraphoxinus (Pisces, Cyprinidae). By E. TREWAVAS .

Cymothoidae) parasitic
By R. J. LINCOLN
: Eunicidae), a member
the western equatorial

Index to Volume 21

67
99

213

257

285
289

297
313
329

339
345

349
359
363

DECAPOD CRUSTACEAN LARVAE

COLLECTED DURING THE
INTERNATIONAL INDIAN OCEAN
EXPEDITION. FAMILIES RANINIDAE
AND HOMOLID

A. L. RICE

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 21 No. i

LONDON : 1970

DECAPOD CRUSTACEAN LARVAE COLLECTED

DURING THE INTERNATIONAL INDIAN
OCEAN EXPEDITION. FAMILIES RANINIDAE

AND HOMOLIDAE

ANTHONY LEONARD RICE

Pp. 1-24; 9 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)

ZOOLOGY Vol. 21 No. i

LONDON: 1970

THE BULLETIN OF THE BRITISH MUSEUM

(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical series.

Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.

In 1965 a separate supplementary series of longer
papers was instituted, numbered serially for each
Department.

This paper is Vol. 21 No. i of the Zoological
series. The abbreviated titles of periodicals cited
follow those of the World List of Scientific Periodicals.

World List abbreviation
Bull. Br. Mus. nat. Hist. (Zool.).

Trustees of the British Museum (Natural History), 1970

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued 22 December, 1970 Price

DECAPOD CRUSTACEAN LARVAE COLLECTED

DURING THE INTERNATIONAL INDIAN

OCEAN EXPEDITION. FAMILIES RANINIDAE

AND HOMOLIDAE

By A. L. RICE

SYNOPSIS

Five crustacean larvae belonging to the family Raninidae and two belonging to the family
Homolidae are described from material collected during the International Indian Ocean exped-
dition. Probable identities of the larvae are suggested where possible. Larval evidence for
the relationship between the Homolidae, the Raninidae and the higher Brachyura is discussed.

INTRODUCTION

THE larval stages of decapod crustaceans of the Indian Ocean are rather poorly
known, few having been hatched from the adults and even fewer reared through all
the larval stages. Consequently, only a very small proportion of larvae taken in
the plankton in this region can be identified to species with any certainty, and in
many cases identification even to family is difficult.

In those decapodan families in which the larval characteristics are already well
known an account of plankton caught material of unknown specific or even generic
identity is of doubtful value. On the other hand, the larvae of some families are so
poorly known that any information on the developmental stages is of value, even if
the material on which this information is based is at the moment unidentifiable.
Such is the case with the crab-like families Raninidae and Homolidae, and although
the I.I.O.E. collections contain very little material of either group a report on it is
warranted.

Family RANINIDAE

Larval stages belonging to some ten species of raninids have been described
previously, but in only three cases have the larvae been definitely identified with a
known adult. Ranina ranina (L.) larvae were hatched from the egg by Aikawa
(1941) and the first stage described. Lyreidus tridentatus de Haan larvae were
reared from the egg to the moult from the 5th to the 6th (last) zoeal stage by William-
son (1965) who also had plankton caught megalopae which moulted in the laboratory
to the first young crab stage. Finally, Knight (1968) reared larvae of Raninoides
benedidi Rathbun taken from the plankton off the Pacific coast of Mexico, some
specimens collected as first zoeae surviving into the early crab stages.

Two of the above species, R. ranina and L. tridentatus, have been recorded as
adults from the Indian Ocean, but the larvae of the other eight recorded Indian

A. L. RICE

TABLE i
Indian Ocean records of adult raninid crabs

Species

Cosmonotus grayi
Adams and White

Lyreidus channeri
Wood-Mason

Lyreidus tridentatus
de Haan

Notopus dorsipes
(Fabr.)

Notosceles chimmonis

Bourne

Notosceles viaderi Ward
Ranina ranina (L.)

Raninoides hendersoni

Chopra
Raninoides personatus

\Vhite

Raninoides serratifrons
Henderson

Locality

Persian Gulf
Dar-es-Salaam
Holothuria Bank,

i335'S: i26E
Bay of Bengal,

21 6'3o"N: 89 2o'E
Bay of Bengal, 9 i4'io"N: 75 46'E
Bay of Bengal
Andaman Sea
"Both sides of Ceylon"
Malabar Coast

Dar-es-Sallaam

N.E. Laurence Marques, Mozambique,

2532'S: 33 2 4 'E
Malabar Coast
Andamans
Zanzibar
Mauritius
Amirante Islands
Seychelle Islands
Mauritius
Durban
Delagoa Bay
Zululand Coast
Mozambique Channel,

I 9 5'S: 36 2i'E
Mozambique
Mauritius
Reunion
Andaman Sea,

n49'5o"N:92 52'E
Bay of Bengal,

off the mouth of the river Hughli
Bay of Bengal
Holothuria Bank
Off Cape Negrais, Burma

i525'N: 9 3 45'E
Off Travancore coast,

955'N: 7545'E
Off southern Ceylon,

6 2'3"N: 81 29'E
Ceylon
Ceylon

Malabar Coast
Port Shepstone, Natal

Source

Alcock, 1896
Doflein, 1904

British Museum (Nat. Hist.

Wood-Mason, 1886
Alcock, 1899
Alcock, 1896
Alcock, 1896
Alcock, 1896
Alcock, 1896

Doflein, 1904

U.S. Nat. Mus.
Alcock, 1896
Alcock, 1896
Nobili, 1905
Studer, 1882
U.S. Nat. Mus.
U.S. Nat. Mus.
Ward, 1942
Barnard, 1950
Barnard, 1950
Barnard, 1950

U.S. Nat. Mus.
Bianconi, 1851
Bouvier, 1915

Hoffman, 1874

Chopra, i933b

Chopra, I933a
Alcock, 1896
Henderson, 1893

Zool. Survey India
Zool. Survey India

Zool. Survey India
Laurie, 1906
Alcock, 1896
Alcock, 1896
Barnard, 1950

INDIAN OCEAN RANINID AND HOMOLID LARVAE

FIG. i. Ranina ranina (L.), stage I zoea. A, Posterior view; B, lateral view; C, frontal view;
D, dorsal view of abdomen; E, enlarged frontal view of lateral carapace spine of left-hand
side; F, enlarged dorsal view of postero-lateral part of telson. Bar scale represents 2.0 mm
for A, i-o mm for B, C and D, and 0-2 mm for E and F.

A. L. RICE

FIG. 2. Ranina ranina (L.), stage I zoea. A, Antennule; B, antenna; C, mandibles; D,
maxillule; E, maxilla; F, first maxilliped; G, second maxilliped. Bar scale represents 0-5 mm.

INDIAN OCEAN RANINID AND HOMOLID LARVAE 7

Ocean raninids (see Table i) are completely unknown. The 1. 1. O.K. collections
contain only four raninid zoeae and a single megalopa. One of these zoeae appar-
ently belongs to R. ranina but the identities of the other specimens are very un-
certain.

The British Museum (Natural History) collections contain hatched material of
R. ranina and I have taken this opportunity to re-illustrate the first zoea of this
species.

Ranina ranina (L.)

(figs i and 2)
Aikawa, 1941, pp. 117-118, fig. i.

MATERIAL: (a) About 100 stage i zoeae hatched in the Aquarium de Noumea,
New Caledonia, 4. XII. 1956 and presented to the British Museum (Natural History)
by Mons. P. Budker of the Museum National d'Histoire Naturelle, Paris (B.M. reg.
no. 1958: 7: 4: 2-9).

(b) One stage i zoea taken by the Anton Bruun at station 18, 07 4i'N: 97 59 'E
on 21. III. 1963. This specimen is badly damaged but appears to belong to this
species.

SIZE: Tip to tip of the rostral and dorsal carapace spines 6-7-7-7 mm -

REMARKS: Because of the relative abundance of material available it is now
possible to illustrate the larvae of this species adequately and a written description is
not necessary.

Aikawa's Japanese larvae were somewhat smaller than those from Noumea and
he gives a spine tip to spine tip length of 4-2 mm, but the two sets of larvae show
very close agreement in all other features which Aikawa either described or illus-
trated.

However, Aikawa did not mention the spinules on the antennae and maxillipeds
although they were almost certainly present in his material. Williamson (1965)
reported such spines on the maxillipeds of first stage zoeae of Lyreidus tridentatus,
and these features may therefore be widespread within the Raninidae. However,
without dissection the appendages of the larvae reported below could not be exam-
ined at sufficiently high magnifications in all cases to determine whether the spinules
are present or not.

Raninid larva A; fRaninoides sp.

(fig- 3)

MATERIAL: One specimen in the first zoeal stage. Position 12 36'N: 8o4o'E;
depth 200 m to surface. Date 20. VI. 1964. Vessel I.N.S. Kistna; Station 378.
I.O.B.C. serial no. 0612.

SIZE: Tip to tip of the rostral and dorsal carapace spines 3-3 mm; Carapace
length from between eyes to postero-lateral carapace margin 0-93 mm.

A. L. RICE

FIG. 3. Raninid larva A, stage I zoea. A, Frontal view; B, lateral view; C, dorsal view of
abdomen; D, antenna; E, first maxilliped ; F, second maxilliped. The setae covering the
surface of the carapace and abdomen are omitted from B and C for clarity. Bar scales repre-
sent i-o mm

INDIAN OCEAN RANINID AND HOMOLID LARVAE g

DESCRIPTION: Carapace with prominent, curved, dorsal and rostral carapace
spines, each with a number of subsidiary spines, mainly on the anterior edge of the
rostral spine and the posterior edge of the dorsal spine. Lateral spines normally
directed downward and forward, roughly parallel to the rostral spine, but that of
the left-hand-side displaced backwards (see fig. 3). Each lateral spine with a row of
subsidiary spines on the dorsal edge. Anterior dorsal tubercle in the mid-line
between eyes; posterior tubercle behind the dorsal spine. A pair of spines on each
side of the posterior carapace tubercle, close to the posterior carapace margin.
Surface of carapace covered with stiff, almost spine-like bristles (not shown in fig.
3(B)). Eyes sessile, each with a proment spine and a papilla on the stalk.

Abdomen of 5 segments plus the telson. Segment i with a large mid-dorsal spine
and 2 pairs of smaller dorso-lateral spines. Segments 2-5 each with a single median
ventral spine and paired dorsal, dorso-lateral, postero-lateral and postero- ventral
spines. The dorso-lateral spines on segment 2 directed forwards like the 'lateral
knobs' on this segment in the larvae of the higher Brachyura.

Telson (fig. 3(C)). Each arm of the shallow telson fork with a very large setose
spine and 3 postero-median plumed setae. Outside the major spine on each side
there are 2 spines, the outer naked, the inner setose. Antero-lateral margin of each
telson arm carries 4 smooth spines. There are a number of spinules on the dorsal
surface of the telson, and each of the 2 major spines has a subsidiary spine on the
dorsal surface close to the base.

Antennule simple and unsegmented, with 2-3 terminal aesthetascs and i seta.

Antenna as shown in fig. 3 (D).

Mandibles with incisor and molar processes, but no palp.

Maxillule with an unsegmented endopod with 3 terminal and i sub-terminal seta.
Endites well developed, but no lateral seta on basis.

Maxilla with 5 setae on the endopod. Scaphognathite with 4 sub-equal plumose
setae on the lateral margin and a much longer posterior seta.

Maxillipeds i and 2 as shown in fig. 3 (E and F).

None of the more posterior appendages developed.

REMARKS: This larva is tentatively attributed to the genus Raninoides, on the
basis of its close resemblance to that of Raninoides benedicti, but it could belong
to other Indian Ocean genera whose larvae are unknown. It differs so much from
the first zoea of Lyreidus tridentatus, particularly in the length of the lateral carapace
spines and in the form of the telson, that it is very unlikely to belong to L. channeri,
the only other member of the genus recorded from the Indian Ocean. The larva also
differs from the Ranina ranina larvae illustrated above, particularly in being much
smaller and possessing eyestalk spines.

Of the raninids recorded from the Indian Ocean this leaves 7 species belonging to
the genera Cosmonotus, Notopus, Notosceles and Raninoides as possible parents.
The larva could belong to any of these genera, but its very close resemblance to the
described larvae of Raninoides benedicti Rathbun (Knight, 1968) makes another
species of this genus a strong possiblity.

The larva differs from the first stage of R. benedicti in only a few relatively minor
points. Thus, the lateral carapace spines are relatively longer than in R. benedicti

A. L. RICE

FIG. 4. Raninid larva B, stage III (?) zoea. A, Frontal view; B, lateral view; C, dorsal view
of abdomen; D, antenna; E, first maxilliped; F, second maxilliped. Bar scale represents
i-o mm for D and 2-0 mm for the remainder.

INDIAN OCEAN RANINID AND HOMOLID LARVAE n

and carry more subsidiary spines. There are 4 spines on the antero-lateral margin
of each telson fork as opposed to 2 in R. benedicti, and there is an extra pair of spines
on the posterior carapace margin. All of these differences are probably specific
rather than generic.

If the larva does belong to Raninoides then the known distributions of the adults
(Table i) indicate that it probably belongs either to R. personatus White or to
R. serratifrons Henderson.

Raninid larva B

(fig. 4)

MATERIAL: One slightly damaged specimen, probably a third zoea. Position
06 26 'N: 49 46'E; depth 200 m to surface. Date 17. VIII. 1964. Vessel Argo;
Dodo cruise; Station 37. I.O.B.C. serial no. 0374.

SIZE: Carapace length from between the eyes to posterior carapace margin
1-83 mm; Abdomen length 2-71 mm; no spine tip to spine tip measurement can be
given because the dorsal spine is broken.

DESCRIPTION: Carapace with long dorsal and rostral spines, and curved, for-
wardly directed lateral spines, each with a number of subsidiary spines. Anterior
and posterior blunt dorsal tubercles in the mid-line. A supra-ocular spine present
on the right-hand-side; the left-hand-side is damaged, but does not appear to have
possessed a spine in this position. A number of spines on the carapace surface
beneath the insertion of each lateral spine and on the slightly raised ridge between
the dorsal and lateral spines. A series of spines on each postero-lateral carapace
margin decreasing in size ventrally. Eyes free, with the stalks carrying many
setae but no sign of papillae or spines. Whole surface of carapace covered with
short setae (not shown in the illustrations).

Abdomen of 6 segments plus the telson. Segments 2-5 with long, acute spines on
the postero-lateral corners. Segment 2 with forwardly directed lateral processes
and segments 4 and 5 each with a median ventral spine. All abdominal segments
with a number of other spines on the dorsal and lateral surfaces and also on the
posterior margins; segments 1-5 have long setae on the posterior margins (fig. 4(C)).

Telson (fig. 4(C)) a broad flat plate, about 4 times as broad as its length in the
mid-line. Posterior margin on the right-hand-side carries a large, fused spine,
presumably representing the 4th telson process. Outside this large spine there are
a very small and 2 larger fused spines which probably represent the first 3 telson
processes, although the outer one is not clearly distinguishable from the series of
spines on the antero-lateral telson margin. Postero-lateral region of the telson on
the left-hand-side is damaged beyond the base of the main fused spine. Posterior
margin with 21 articulated processes between the major spines. A series of small
spines on the dorsal surface, close to the posterior margin, extends onto the main
telson spines. Dorsal surface of telson and abdominal segments with many small
setae.

A. L. RICE

Antennule simple and unsegmented, with 3-4 terminal aesthetascs and a single
seta.

Antenna (fig. 4(D)) with 19-20 marginal setae on the scale and a series of small
bristles on both the upper and lower surfaces. Endopod less than 1/3 length of

FIG. 5. Raninid larva C, terminal zoea. A, Dorsal view; B, lateral view of thorax; C, lateral
view of abdomen; D, antenna; E, ventral view of last pleopod and uropod on the right-hand-
side. The small surface setae have been omitted partly from A and wholly from B. Bar
scales represent i-o mm.

INDIAN OCEAN RANINID AND HOMOLID LARVAE 13

scale, fused to basis and armed with a single terminal seta. Basis with spinous
process about 1/3 length of scale, and a second prominent spine at the base of the
scale.

Manbidle with no palp.

Maxillule with an unsegmented endopod armed with 4 setae. A single seta on
the lateral margin of the basis below the endopod.

Maxillipeds i and 2 as shown in fig. 4(E and F). Exopod of maxilliped i with n
or 12 natatory setae, that of maxilliped 2 with 13 setae on the distal half and 3 on
the proximal half.

Maxilliped 3 and the more posterior thoracic appendages present as unarmed,
unsegmented buds.

No pleopods present but the uropods have well developed exopods carrying 6 or 7
marginal setae. Endopods represented by very small unarmed buds not separated
from the protopods.

REMARKS : From the known development of Lyreidus tridentatus and Raninoides
benedicti it is clear that the determination of the zoeal stage in raninid larvae is not
as simple as it is, for example, in the higher Brachyura. However, from the pres-
ence in the above larva of well developed uropods but the absence of pleopods, and
also the degree of development of the antennules, antennae and maxillipeds, it
appears to be at about the mid-point of the zoeal series, perhaps the third or fourth
of a total of about 6 zoeal stages.

Identification of this larva is also difficult. The differences between it and the
known larvae of Lyreidus and Raninoides suggest that it belongs to a genus other
than these. The changes which would be necessary to transform the first zoea of
Ranina ranina into this larva, particularly in the armature of the carapace and the
form of the telson, are greater than those ocurring during the development of L.
tridentatus and R. benedicti, indicating that it does not belong to R, ranina either.
This leaves the parentage still very uncertain, but probably among the genera
Cosmonotus, Notopus and Notosceles.

Raninid larva C

(fig- 5)

MATERIAL: One specimen in the last zoeal stage. Position 09 34'N: 75 i6'E;
depth 200 m to surface. Date 10. II. 1965. Vessel Meteor; Station 187. I.O.B.C.
serial no. 0145.

SIZE: Tip of rostral spine to tip of dorsal spine 3-8 mm; tip of rostral spine to
posterior median carapace margin 3-9 mm; total length (tip of rostral spine to base
of telson fork) 6-6 mm.

DESCRIPTION: Carapace with prominent forwardly directed rostral and paired
lateral spines; subsidiary spines on the dorsal surface of the rostrum and on the
outer surfaces of the laterals. Dorsal spine small, curving posteriorly. Blunt
anterior and posterior carapace tubercles in the mid-line. Rostrum widens between
the eyes to a front carrying a series of spines on each antero-lateral angle. A pair

14 A. L. RICE

of bulbous lobes behind the eyes each carrying a prominent spine antero-laterally
and a number of smaller spines dorsally. A curving row of 6 spines on each side
of the carapace between the bases of the lateral and dorsal spines. A parallel row of
4 spines close to the postero-lateral carapace margin. Each postero-ventral corner
of the carapace with a single spine and a series of denticles. A prominent papilla on
each eyestalk.

Abdomen (fig. 5 (A and C)) of 6 segments plus the telson. First segment with a
raised transverse ridge armed with a row of long plumose setae and with 3 spines
close to the mid-line. Segments 2-5 all basically similar, having each postero-
lateral margin produced into a long, slightly curved spine. A somewhat variable
number of other spines on each segment, but a basic pattern of 4 on the dorsal
surface anteriorly, and a series of 5 on the postero-dorsal margin. Segment 5 with
additional spines, including a prominent one at each postero-lateral corner. Seg-
ment 6 with only 2 dorsal spines, but 6 spines on the posterior margin, the outer
pair in dorsal view looking like the postero-lateral spines of the more anterior seg-
ments.

Telson (fig. 5 (A)) a broad, bilobed plate. Each antero- and postero-lateral margin
with a series of fused spines increasing in size gradually posteriorly, but ending in a
much larger spine with a small spine basally on the dorsal surface. Posterior telson
margin with 10 pairs of articulated processes between the telson forks, inner 3 pairs
much smaller than the others and probably added at the most recent moult.

The whole of the dorsal surface of the carapace, abdomen and telson covered with
short, close-set setae.

Antennule with an unsegmented peduncle, swollen basally and with an obvious
statocyst. Endopod unarmed and not separated from the peduncle. Flagellum
with 3 groups of about 3, 3, and 4 aesthetascs.

Antenna (fig. 5(D)) with 26-28 marginal setae on the scale. Endopod 3-segmented,
about as long as scale and with a single terminal seta. Protopod with 2 spines and

3 setae.

Mandible with an unarmed, unsegmented palp.

Maxillule with 2-segmented endopod carrying 5 setae on the distal segment and a
single seta on the proximal segment. Lateral margin of basis with a single seta.

Maxilla with more than 70 marginal setae on the scaphognathite. Unsegmented
endopod with 6 terminal, i medial and 4 lateral setae.

Maxilliped i with 2-lobed epipod. Basipod with 15 setae on medial margin.
The 5 segments of the endopod carry 3, 2, 2, 4 and 6 setae respectively on the medial
margins. The lateral margins of segments 2 and 5 (terminal) each with a single
fine seta, those of segments 3 and 4 each with 2 fine setae. Exopod 2-segmented;
proximal segment with 8 setae along posterior edge, distal segment with 15 or 17
setae around whole margin.

Maxilliped 2 with simple epipod. Basipod with 4 setae on medial margin and 2 on
posterior surface at the base of the endopod and exopod. Endopod on one side of

4 segments with 2, i, 2 and 3 inner setae respectively. On the other side the long
penultimate segment is sub-divided, the proximal part being unarmed. Terminal
and sub-terminal segments each with a very fine lateral seta. Exopod 2-segmented ;

INDIAN OCEAN RANINID AND HOMOLID LARVAE 15

proximal segment with 6 or 7 setae along posterior edge, distal segment with 21
marginal setae.

Maxilliped 3 with endopod indistinctly divided into 4 segments of which the
terminal and sub-terminal each carry a short seta. Exopod short, simple and
unarmed.

Pereiopods all present and indistinctly segmented, the first pair chelate.

Pleopods on abdominal segments 2-5 well developed but unsegmented, with small
endopods and larger exopods (fig. 5 (C and E)). Exopods and endopods unarmed,
but with somewhat serrate margins indicating that they would probably become
setose at the next moult. Uropods with exopods separated from protopods and
carrying 20-21 plumose setae. Endopods represented by small buds and not
separated from the protopods (fig. 5(E)).

REMARKS: Of the raninid larvae previously described, this zoea most closely
resembles Lithozoea serrulata described by Aikawa (1933). The two larvae share
a number of common features including the broad, plate-like telson, forwardly
directed carapace spines, spine rows on the posterior carapace margin and on the
side of the carapace between the lateral spines and the dorsal spine, and relatively
short rostral and dorsal spines. The main differences between the larvae are the
development of the anterior carapace tubercle into a bifurcated spine and the pres-
ence of lateral carapace keels in Lithozoea, the presence of a mandibular palp in the
Indian Ocean larva and 4 ot 5 segments in the endopod of the second maxilliped
in this larva compared with only 3 in Lithozoea.

Williamson (1965) pointed out that Lithozoea serrulata possesses a number of
homolid characters, particularly in the carapace spines and keels. Although raninid
larva C is somewhat less homolid in these respects than Lithozoea, it does resemble
late homolid zoeae in having more than 3 segments in the endopod of the second
maxilliped. No brachygnathan larvae have more than 3 segments in this endopod,
and the only previously described raninid larva which may have more than 3 is
Acanthocaris described by Claus (1876 and 1885) (see Williamson, 1965, p. 388),
which also shows some homolid features in its carapace.

Little can be said about the identity of this Indian Ocean larva. As with the
previous larva, comparison with published larval descriptions readily excludes it
from the genera Lyreidus, Ranina and Raninoides. This still leaves as possible
parents the genera Notopus, Cosmonotus and Notosceles, although adult Notosceles
are so similar to Raninoides that they must surely have similar larvae. The simil-
arities between the Indian Ocean larva and Aikawa's Lithozoea serrulata suggest that
they belong to the same genus. If this is so, the Indian Ocean larva may belong to
Notopus dorsipes (Fabr.) since, of the three genera deduced as possible parents,
Notopus is the only one with different species recorded in Indian and Japanese
waters. N. dorsipes is the only species known from the Indian Ocean, having been
recorded by Alcock (1896) from off the Malabar coast, relatively close to where
raninid larva C was collected, but two other species, N. ovalis Henderson and N.
misakiensis Sakai are also recorded from Japan (Sakai, 1937). Lithozoea serrulata
perhaps belongs to one of these species.

A. L. RICE

Raninid larva D; fRaninoides sp.

(figs 6 and 7)

MATERIAL: One megalopa. Position 10 36'N: 95 39'E; depth 125 to 250 m.
Date 25. III. 1963. Vessel Anton Bruun; cruise i; Station 24 S.O.S.C. Ace. no. 3.

SIZE : Carapace length from tip of rostrum to posterior carapace margin 3 -6 mm ;
Maximum carapace width 1-9 mm.

DESCRIPTION: The megalopa is fully illustrated in figs 6 and 7. It is very
similar to the megalopa of Raninoides benedicti as described by Knight (1968) and
therefore only the main differences between the two larvae will be mentioned here.

FIG. 6. Raninid larva D; ? Raninoides megalopa. A, Dorsal view; B, lateral view of cara-
pace; C, ventral view of anterior part of thorax; D, dorsal view of telson; E, cheliped, left-
hand-side, ventral view; F, G, and H, third, fourth and fifth pereiopods respectively of the
right-hand-side, dorsal views. Bar scales represent i-o mm for A, B, C and E, and 0-5 mm
for D, F, G and H.

INDIAN OCEAN RANINID AND HOMOLID LARVAE

FIG. 7. Raninid larva D ; ?Raninoides megalopa. A, Maxillule; B, maxilla; C, D and E, first,
second and third maxillipeds in ventral view; F, ischium and exopod of third maxilliped in
dorsal view. Bar scale represents i-o mm.

i8 A. L. RICE

The carapace is slightly narrower posteriorly than is that of R. benedicti. The
eyestalk papillae are placed postero-ventrally rather than antero-dorsally as in R.
benedicti. The large spine between the bases of the chelipeds in R. benedicti is
absent. The Indian Ocean megalopa has no podobranch on the second maxilliped,
while the ischium of the third maxilliped (fig. y(E and F)) is relatively longer than in
R. benedicti and has no teeth on the inner margin.

The chelipeds and the second and third legs in R. benedicti each have a prominent
spine on the ventral margin of the merus; the Indian Ocean megalopa lacks these
spines but has a spine on the dorsal margin of the carpus of legs two and three.
(These two legs are very similar, and therefore only leg three is illustrated in fig. 6.)
The fourth and fifth legs are very similar in shape in the two larvae, but the fourth
leg in the Indian Ocean specimen carries many more setae and the dactyl is less
acutely pointed than in R. benedicti.

REMARKS: The similarity between this larva and the known megalopa of Ran-
inoides benedicti indicates that it belongs to the same genus. From the locality
of capture of the larva and the known distributions of the adults, Raninoides hender-
soni Chopra and R. serratifrons Henderson are equally likely to be the parent.

Family HOMOLIDAE

Plankton-caught larvae of the Homolidae present very much the same problems
of identification as do those of the Raninidae, since larvae have been hatched from
only four species and the complete larval development is known for only one of these
(see Williamson, 1965 and Rice and Provenzano, 1970).

The 1. 1. O.K. collections contain only two homolid zoeae, neither of them agreeing
with any previously described larvae.

Homolid larva A fHomola sp.

(fig. 8)

MATERIAL: One specimen, probably a second zoea. Position 11 49 'S: 49 23 'E.
Date 21. VII. 1964. Vessel R.R.S. Discovery; Station 5508. I.O.B.C. serial no.

SIZE : Carapace length from tip of rostrum to posterior margin in mid-line i -4 mm ;
Total length from tip of rostrum to posterior margin of telson in mid-line 2-5 mm.

DESCRIPTION : The main features of the morphology of this larva are adequately
shown in the illustrations. This description will therefore be restricted to those
features which are not illustrated but which can be seen without dissection.

Antennule 2-segmented. Distal segment with 3 or 4 aesthetascs and setae;
proximal segment with a long median seta, representing the inner flagellum, and a
row of 3 short setae.

Mandibles without palps.

Maxilla with about 18 marginal setae on the scaphognathite.

Maxilliped I as shown in fig. 8 (D).

INDIAN OCEAN RANINID AND HOMOLID LARVAE 19

Maxilliped 2 with the 4 segments of the endopod carrying i, i, 2 and 5 setae
respectively, one seta on the terminal segment being laterally placed. Basis with 5
medial setae and the exopod with 7 natatory setae.

Maxilliped: j with an unsegmented endopod carrying a single terminal seta.
Exopod with 7 setae.

Posterior thoracic appendages represented by unarmed, unsegmented buds.

FIG. 8. Homolid larva A, stage II zoea. A, Dorsal view of carapace; B, dorsal view of
abdomen; C, lateral view; D, antenna; E, first maxilliped. Bar scale represents i-o mm.

20 A. L. RICE

REMARKS: In general form this larva is very similar to the larvae of Homola
barbata (Fabr.) described from Florida by Rice and Provenzano (1970), and also
to the series of larvae taken in the plankton off South Africa and attributed to a
species of Homola by Rice and von Levetzow (1967). However, the present specimen
differs from these previously described forms in several important respects.

The supra-ocular spines and the anterior and posterior carapace tubercles, which
are very prominent in the Florida and South African larvae, are much reduced in the
Indian Ocean specimen. This larva also has the anterior end of the dorsal tooth
row on the carapace less protruberant than in the other larvae, and less likely to be
replaced by a spine in the later stages.

The dorsal spines on abdominal segments 2-5 in the Florida and South African
larvae are entirely lacking in this specimen and, except for the second segment, the
dorso-lateral spines are reduced to blunt protruberances. The Indian Ocean larva
also lacks spines on the dorsal surface of the telson.

Finally, in its degree of development this larva does not agree precisely with any
stage in either of the previously described series, falling between the second and
third stage in both cases.

The absence of dorsal telson spines in two 1 Homola larvae from off south-east
Africa described by Boas (1880) suggested to Rice and von Levetzow that these
larvae were specifically distinct from their own South African material. From the
known distributions of the adults these authors suggested that Boas's larvae might
represent Homola orientalis Henderson, while their own material belonged to the
eastern Atlantic and South African form of H. barbata.

The larvae described here also lacks dorsal telson spines, like Boas's larvae, but
differs from them in having reduced supra-ocular spines. Boas's larvae also
have a pair of dorso-lateral carapace spines, presumably developed from the anterior
end of the dorsal carapace tooth row of an earlier stage ; as noted above, such spines
are unlikely to be developed in a later stage of the I.I.O.E. larva. If these differences
are specific, then there is either a third, unsuspected, species of Homola in the south-
western Indian Ocean, or else at least some of the larvae attributed to Homola in
fact belong to a different genus.

Knowledge of generic differences between homolid larvae is restricted to a compari-
son of hatched larvae of Homola barbata (Fabr.) (Rice and Provenzano, 1970),
Paromola japonica Parisi and Latreillia phalangium de Haan (Aikawa, 1937) and
Latreillia australiensis Henderson (Williamson, 1965). Williamson noted that the
homolid larvae described up to that time (1965) fell into two groups, one of which,
including the hatched larvae of Latreillia and Paromola, differed from the second
group in having neither dorsal nor antero-lateral carapace spines and no dorsal
spines on the abdomen. Williamson suggested that these groups probably did not
represent taxonomic groupings within the family and that intermediate forms would
probably be found as more homolid larvae were described.

The larva described here agrees most closely with the second of these larval
groups, which includes not only the larvae described by Rice and von Levetzow
(1967), Rice and Provenzano (1970) and Boas (1880), but also Gurney's (1924)
Dromiacean species I and specimens attributed to the genus Homola by Cano (1893),

INDIAN OCEAN RANINID AND HOMOLID LARVAE

Thiele (1905), Pike and Williamson (1960) and Rice (1964). However, it differs
from all of these forms in the absence of mid-dorsal spines on the abdominal segments
and is therefore intermediate between Williamson's two groups. Considering the
great similarity between the known larvae of Latreillia and Paromola, generic
differences between homolid larvae seem to be very slight so that this larva may,
indeed, belong to a genus other than Homola.

FIG. 9. Homolid larva B, stage III zoea. A, Dorsal view; B, lateral view; C, antenna; D, E
and F, first, second and third maxillipeds. Bar scale represents i-o mm for A and B, and
0-5 mm for C, D, E and F.

22 A. L. RICE

Homolid larva B

(fig- 9)

MATERIAL: One specimen, probably a third zoea. Position 16 3i'N: 54 o8'E.
Date 30. VI. 1963. Vessel R.R.S. Discovery: Station 5026. I.O.B.C. serial no. 0811.

SIZE : Carapace length from tip of rostrum to posterior margin in mid-line i -5 mm.
Total length from tip of rostrum to posterior margin of telson in mid-line
3 -3 mm.

DESCRIPTION: Carapace with a forwardly directed and somewhat upturned
rostrum. Prominent mid-dorsal spine and smaller posterior carapace tubercle; no
anterior tubercle. Dorsal tooth row of 70-80 teeth on each side, extending from
middle of posterior margin to behind the eye and ending in a rather longer and
stouter tooth. Postero-ventral tooth row of 50-60 sub-equal teeth. A prominent
ventro-lateral spine on each side of the carapace above the base of the third maxil-
liped. Prominent eyestalk papillae.

Abdomen of 6 segments and telson. Segment 2 with slight dorsal transverse ridge
and segment 5 with small postero-lateral processes; otherwise all abdominal seg-
ments simple and unarmed apart from some setae close to posterior margins.

Telson a broad, triangular plate, with 3 fused spines at each postero-lateral angle
and 10 pairs of processes articulated to posterior margin. Uropods with protopods
not separated from endopods and exopods. Endopods unarmed, exopods with
9-11 marginal setae.

Antennule 2-segmented, proximal segment with a single long plumose seta repre-
senting ventral flagellum, distal segment with 2 terminal aesthetascs and 2 setae.

Antenna (fig. g(C)) with 16 marginal setae on scale, spinous process about 1-5 times
length of scale, endopod unarmed and less than half length of scale.
Mandibles without palps.

Maxillule with 5 setae on distal segment of endopod and a single seta on proximal
segment ; no lateral seta on basis.

Maxilla with about 40 setae on scaphognathite.

Maxillipeds as illustrated in figs 9 (D, E, and F) ; exopods of each with 8 natatory
setae.

Posterior thoracic appendages present as unarmed, unsegmented buds.

REMARKS: Like the preceding larva, this specimen is intermediate between the
two homolid larval groups noted by Williamson. Thus, although it possesses the
prominent dorsal carapace spine of the 'Homola group, it lacks the dorsal abdom-
inal spines typical of most of these larvae. It is even less Homola-like. than the larva
described above since it also lacks any suggestion of supra-ocular spines, it has very
small antero-laterals (if, indeed, the enlarged teeth at the anterior end of the dorsal
carapace tooth rows represent these), and it has no dorso-lateral projections on the
abdominal segments. The combination of characters is so different in this larva
from that in any previously described zoea that its identity cannot be established at
present.

INDIAN OCEAN RANINID AND HOMOLID LARVAE 23

DISCUSSION

The small collection of raninid and homolid larvae reported here tends to confirm
Williamson's (1965) conclusions that the similarities between the two groups are
consistent with both families being fairly close to a pre-brachyuran stock, the
raninids being considerably more 'brachyuran' than the homolids.

Perhaps the most interesting larva in the collection is that described as raninid
larva C which, like Aikawa's Lithozoea serrulata, is intermediate in many features
between the homolids and the typical raninid larvae. Williamson noted that L.
serrulata represents a possible intermediate step in the simplification of the rather
complex carapace armature of the homolids towards the dorsal, rostral and lateral
spines of the carapace in typical raninid and brachygnathan zoeae. Raninid larva C
seems to be a further step in this direction, since the homolid denticulate carapace-
folds, still present in L. serrulata, are represented only by a series of short spines in
this larva. The relationship of the lateral carapace spines to the carapace folds and
denticle rows in raninid larva C and L. serrulata indicate that the lateral spines in the
raninids, and presumably also in the higher Brachyura, are homologous with either
the antero-lateral spines in the homolids or, more probably, with the spines developed
from the anterior ends of the dorsal denticle rows.

The bilobed, plate-like telsons of L. serrulata and raninid larva C also conveniently
bridge the gap between the rather anomuran broad triangular telsons of late homolid
zoeae and the forked telsons of the zoeae of Ranina, Raninoides and Lyreidus, which
are more similar to those of typical brachyuran zoeae.

It would be interesting to know if these seemingly primitive raninid larvae belong
to species with similarly primitive adults, but such information can come only from
the rearing work which is much needed in both the Homolidae and the Raninidae.

ACKNOWLEDGEMENTS

My thanks are due to the Directors of the Indian Ocean Biological Centre and the
Smithsonian Sorting Center for allowing me to examine material in their care. I
would also like to thank Dr. K. K. Tiwari of the Zoological Survey of India and Dr.
Henry B. Roberts of the Smithsonian Institution for providing me with details of
adult raninids from the Indian Ocean in their collections.

REFERENCES

AIKAWA, H. 1933. On larval forms of some Brachyura. Paper II: a note on indeterminate

zoeas. Rec. Oceanogr. Wks. Jap. 5 : 124-254.
1937. Further notes on brachyuran larvae. Rec. Oceanogr. Wks. Jap. 9 : 87-162.

- 1941. Additional notes on brachyuran larvae. Rec. Oceanogr. Wks. Jap. 12 : 117-120.
ALCOCK, A. W. 1896. Materials for a carcinological fauna of India. No. 2. The Brachyura

Oxystomata. /. Asiat. Soc. Beng. 65 (II) : 134-296.

- 1899. An account of the deep-sea Brachyura collected by the Royal Indian Museum
Maritime Survey Ship "Investigator". Calcutta, 85 pp.

BARNARD, K. H. 1950. Descriptive catalogue of South African decapod Crustacea (Crabs and

shrimps). Ann. S. Afr. Mus. 38 : 1-837.
BIANCONI, G. G. 1851. Specimina Zoologica Mosambicana. Fasc. 5 : Memorie R. Accad. Sci.

1st. Cl. Sci.fis. Bologna, 3 : 91-112.

24 A. L. RICE

BOAS, J. E. V. 1880. Studier over Decapodernes Slaegskabsforhokl. K. danske Vidensk.

Selsk. Skr. (6) Nat. og Math., Afd. 1 (2) : 25-210.
BOUVIER, E. L. 1915. Decapodes marcheurs (Reptantia) et Stomatopodes recueillis a 1'ile

Maurice par M. Paul Carie. Bull, scient. Fr. Belg. (7) 48 (3) : 178-318.
CANO, G. 1893. Svillupo del Dromidei. Atti Accad. Sci. fis. mat., Napoli 6 (2) : 1-23.
CHOPRA, B. i933a. Further notes on Crustacea Decapoda in the Indian Museum. III. On

the decapod Crustacea collected by the Bengal Pilot Service off the mouth of the River

Hughli. Dromiacea and Oxystomata. Rec. Ind. Mus. 35 : 25-52.
- !933b. Further notes on crustacean Decapoda in the Indian Museum. IV. On two new

species of oxystomous crabs from the Bay of Bengal. Rec. Ind. Mus. 35 : 77-86.
CLAUS, C. 1876. Untersuchungen zur Erforschung der genealogischen Grundlage des Crustaceen-

Systems. Ein Beitrage zur Descendenzlehre. VVein, 114 pp.
CLAUS, C. 1885. Neue Beitrage zur Morphologic der Crustaceen. Arb. zool. Inst. Wien, 6 :

1-108.

DOFLEIN, F. 1904. Brachyura. Ergebn. dt. Tiefsee-Expedn 'Valdivia' 1898-1899. 6 :i-3i4-
GURNEY, R. 1924. Decapod larvae. Nat. Hist. Rep. Br. Antarct. Terra Nova Exped. (Zool.)

8 : 37-202.
HENDERSON, I. R. 1893. A contribution to Indian carcinology. Trans. Linn. Soc. Lond. (2)

5 : 325-458.
HOFFMANN, C. K. 1874. Crustaces et Ecinodermes. In: F.P.L. Pollen and D.C. Van Dam

(Ed.) Recherches sur la faune de Madagascar 5 (2) : 1-58.
KNIGHT, M. D. 1968. The larval development of Raninoides benedicti Rathbun (Brachyura,

Raninidae), with notes on the Pacific records of Raninoides laevis (Latreille). Crustaceana,

Suppl. 2 : 145-169.
LAURIE, R. D. 1906. Report on the Brachyura collected by Prof. Herdman at Ceylon 1902.

Rep. Pearl Oyster Fish. 5, Suppl. Rep. XL: 349-432.
NOBILI, G. 1905. Crostacei di Zanzibar. Boll. Mus. Zool. Anat. comp. R. Univ. Torino. 20

(506) : 1-12.
PIKE, R. B. and WILLIAMSON, D. I. 1960. Larvae of decapod Crustacea of the families

Dromiidae and Homolidae from the Bay of Naples. Pubbl. Staz. zool. Napoli 31 : 553-563.
RICE, A. L. 1964. The metamorphosis of a species of Homola (Crustacea, Decapoda: Drom-
iacea). Bull. mar. Sci. Gulf Caribb. 14 : 221-238.

and PROVENZANO, A. J. 1970. The larval stages of Homola barbata (Fabricius) (Crus-
tacea, Decapoda, Homolidae) reared in the laboratory. Bull. mar. Sci. (20 : 446-471).

and VON LEVETZOW, K. G. 1,967. Larvae of Homola (Crustacea, Dromiacea) from South
Africa. /. nat. Hist. 1 : 435-453.

SAKAI, T. 1937. Studies on the crabs of Japan. II. Oxystomata. Scient. Rep. Tokyo

Bunrika Daigaku Sect. B 3, Suppl. no. 2 : 67-192.
THIELE, J. 1905. Uber einige stielaugige Krebse von Messina. Zool. Jb., Suppl. 8 (Festschr.

f. Mobius) : 443-474.
WARD, M. 1942. A new genus and eight new species of Brachyura from Mauritius and the

Chagos Archipelago. Bull. Maurit. Inst. 11 (2) : 39-48.
WILLIAMSON, D. I. 1965. Some larval stages of three Australian crabs belonging to the

families Homolidae and Raninidae, and observations on the affinities of these families

(Crustacea: Decapoda). Aust. J. Mar. Freshw. Res. 16 : 369-398.
WOOD-MASON, J. 1886. Description of a new species of Crustacea belonging to the brachy-

urous family Raninidae. /. Asiat. Soc. Beng. 56 : 206-209.

Dr. A. L. RICE,

Department of Zoology

BRITISH MUSEUM (NATURAL HISTORY)

CROMWELL ROAD

LONDON, S.W.7

Printed in England by Staples Printers Limited at their Kettering, Northants, establishment

BATS FROM ETHIOPIA COLLECTED
BY THE GREAT ABBAI EXPEDITION,

1968

J. E. HILL

AND

P. MORRIS

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 21 No. 2

LONDON: 1971

I 2 7 APR 1971

BATS FROM ETHIOPIA COLLECTED BY
THE GREAT ABBAI EXPEDITION, 1968

JOHN EDWARDS HILL

AND

K
PATRICK MORRIS

Pp. 25-49; 3 Plates

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)

ZOOLOGY Vol. 21 No. 2

LONDON : 1971

THE BULLETIN OF THE BRITISH MUSEUM

(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical series.

Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.

In 1965 a separate supplementary series of longer
papers was instituted, numbered serially for each
Department.

This paper is Vol. 21, No. 2 of the Zoological series.
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Issued 28 April, 1971 Price 1-15

BATS FROM ETHIOPIA COLLECTED BY
THE GREAT ABBAI EXPEDITION, 1968

By J. E. HILL & P. MORRIS

INTRODUCTION

THERE has been hitherto no great concentration on the bat fauna of Ethiopia by
collectors and many records are of one or two specimens sporadically obtained.
For this reason the present collection is of particular interest, and is of especial
value for the wide representation of the bats of western Ethiopia, hitherto largely
unrecorded. This paper is an account of the bats collected by the Great Abbai
Expedition including taxonomic notes, and revisions where appropriate, together
with what little ecological information is available.

The principal aim of the Expedition was to carry out a survey of the Blue Nile
( Great Abbai) Gorge. The organization of the Expedition and its main activities
are described by Blashford Snell (1970). The bat collections were made mainly in
the Gorge itself at river level at various "Forward Bases" west of Shafartak Bridge,
or in the Awash National Park, with some additional specimens being obtained
elsewhere, notably in the highlands at Ghimbi.

The Nile Gorge is a unique habitat, mostly uninhabited with dense scrub covering
the steep rocky hillsides. There is a spectacular seasonal change here; during the
dry season the river almost ceases to flow and the Gorge becomes exceedingly arid
with only a narrow fringe of tall trees beside the river retaining their leaves. At
this time survival must be a serious problem for many animals, Megachiroptera
especially being unlikely to find any succulent fruits and presumably migrating
elsewhere. By contrast insects appear to be especially abundant in the dry season,
according to previous explorers, so that the Microchiroptera may well remain
throughout the year, though the fierce heat and lack of shelter must cause problems.
The Expedition's visit to the Nile Gorge came towards the end of the wet season,
when the atmosphere was warm and humid; the lush green vegetation seemed to
provide an ideal habitat for a wide range of animals and gave no indication of just
how inhospitable the habitat is during at least one third of the year.

The Awash National Park is chiefly an open dusty area of grassland, thorn scrub
and volcanic rubble. An important geological feature, so far as bats are concerned,
is the occurrence of "lava blister caves". These are evidently formed by gas
bubbles in flowing lava and appear on the ground as low, rounded hillocks standing
up prominently in an otherwise flat area. These mounds are hollow inside and
some have perforated so allowing access to the interior. They represent the only
cool, shady retreats in an otherwise open, arid landscape and consequently house
many bats. Two of the blisters had openings only at their highest points. These
had functioned as pitfall traps and also perhaps as carnivore dens, and the floors of
both were littered with mammal bones which included the remains of five bat
species.

28 J. E. HILL & P. MORRIS

Live bats were collected either by shooting (with -22 or -410 guns loaded with
dust shot) or with mist nets. Fixed nets, supported on poles, trees or convenient
rocks were avoided by flying bats, and only proved useful when set at a roost entrance
whence the bats could be driven into the net. For catching bats as they flew
low hawking for food a hand held net was employed. A small net (about i m x
1-5 m), supported between two fibreglass rods was held close to the ground and
flicked sharply upwards to catch the bat as it flew past. The method requires both
patience and a certain amount of practice and was used with particular success by
Dr. D. W. Yalden to catch most ol the free-flying bats obtained. Specimens taken
this way come in ones and twos and contrast with sizeable batches of bats taken at
roosts. Because roosting bats are easier to get in quantity, they figure prominently
in this collection, though their numbers here are not necessarily a reflection of true
abundance in the field. Netted bats are often of greater interest purely because
they are difficult to catch, particularly if they do not form communal roosts ; they
are therefore poorly represented in collections.

The specimens obtained were either preserved whole in formalin, supported on
card so as to keep the wings well displayed or they were prepared as a skull and dry
card-mounted skin. The collections of the British Museum (Natural History)
contain also a small number of specimens representing species previously not recorded
from Ethiopia and notes on these are included in this report. Such specimens are
denoted by their registration numbers, as is a single specimen from French Somali-
land (Territory Afars and Issas) which constitutes a new record for that country and
is included in this paper.

The majority of bat specimens collected by the Great Abbai Expedition are now
in the collections of the British Museum (Natural History), but where numbers have
permitted this to be done, duplicate examples have been sent to the Museum at
Haile Selassie I University, Addis Ababa.

In the present paper, the altitudes and co-ordinates used have been taken from
the best available maps, the i = 1,000,000 series. For convenience, clarity and
ease of future reference these have been adhered to, even though investigations in
the field showed the maps to be inaccurate in places. Part of the Nile forms the
boundary between the provinces of Go j jam and Wollega; specimens obtained at
the riverside have not been specifically assigned to either. Times given are 'local
time', and as a rough rule the period 19.00 hrs-o6.oo hrs is passed in darkness.
Linear measurements of specimens are in millimetres : the minimum, maximum and
mean (in parentheses) are given for series.

It is hoped that a future paper will give detailed background information on the
ecology of the areas visited by the Expedition, meanwhile the brief survey above
and the "collection and field notes" for the various species below have been prepared
by one of us (P.M.) from actual field data; determinations and taxonomic investiga-
tions are the work of J.E.H. We would like to express our thanks to various
members of the Great Abbai Expedition for the hard work and long hours spent
in pursuit of bats under trying conditions. The efforts in the field of Drs. D. W.
Yalden, M. J. Largen and Mr. H. King are particularly acknowledged.

BATS FROM ETHIOPIA 29

SYSTEMATIC SECTION

Epomophorus anurus (Heuglin, 1864)

SPECIMENS. 2 young adult females. Temporary base, Mouth of Azir River,
Blue Nile Gorge, 10 29' N, 36 25' E, alt. 1,000 m. 21 August 1968.

TAXONOMIC NOTES. These young adult specimens agree closely in palatal propor-
tions with those given by Anderson (1912 : 533) for females of E. anurus. Kock
(1969 : 18) considers anurus a subspecies of E. labiatus.

COLLECTION AND FIELD NOTES. A small group of fruit bats was located, flying
about under the big trees lining the river; the bats were particularly concentrated
around some large fig trees which bore plenty of ripe fruit. The remains of many
chewed figs littered the ground below. The bats were seen at about 21.30 hours,
and were immediately recognized as Megachiroptera by the distinctive way in which
their eyes appeared large and red as they reflected the light from torch beams. Three
specimens were obtained using dust shot. It was interesting to note a form of
communal behaviour ; the bats had been following each other in threes and fours
through gaps in the foliage, and when each specimen was shot, the others changed
course and flew low and close to investigate the victim. The bats showed particular
concern in response to the cries of a wounded individual. No other instances of
group behaviour of this nature were observed with any of the other bats obtained.
This communal response is all the more remarkable if the group really did include
two species (see below). All three specimens of Epomophorus were females, so it
may be that the group was part of a nursing colony whose individuals would perhaps
have a stronger attachment to others of their species than normal.

Epomophorus sp.

SPECIMEN, i young adult female. Temporary Base, mouth of Azir River, Blue
Nile Gorge, 10 29' N, 36 25' E, alt. 1,000 m. 21 August 1968.

TAXONOMIC NOTES. It seems likely that this young adult specimen is referable
to E. gambianus, a species recorded from southern Ethiopia by Andersen (1912 :
540). Although from the proportions of the palate it could be referred to
E. crypturus as defined by that author, this species is known so far only from localities
in and south of the southern Congo and southern Tanzania, and, furthermore, in
several of its dimensions this specimen exceeds the greatest size as yet recorded for
E. crypturus. Measurements: length of forearm 77-9; greatest length of skull 50-3;
condylobasal length 50-0; condylocanine length 48-3; median palatal length 27-7;
post palatal length 12-3; rostral length 19-7; zygomatic width - ; least inter-
orbital width 7-5; post orbital width 9-4; width of braincase 16-8; mastoid width
17-2; ci-c 1 (alveoli) 8-9; mi-m 1 13-6; c-m 1 18-2; length of mandible 39-3;
c-m 2 19-6.

COLLECTION AND FIELD NOTES. As for E. anurus. If the present specimen really
is a different species, it is surprising that it formed part of such a closely knit
social group with E. anurus.

30 J. E. HILL & P. MORRIS

Micropteropus pusillus (Peters, 1868)

SPECIMENS. Two females and a foetus. Sirba, Blue Nile Gorge, ioo5'N,
35 30' E, alt. 800 m. 30 August 1968.

TAXONOMIC NOTES. There appears to have been hitherto no confirmed record of
Micropteropus from Ethiopia, although M. pusillus occurs in the southern Sudan
(Kock, 1969 : 24).

COLLECTION AND FIELD NOTES. Found at night between 20.00 hrs and 21.00 hrs
flying around under large fig trees at the edge of the Nile. Both specimens caught
in small hand-held mist nets. One animal was found to be pregnant with a well
formed foetus, suggesting that breeding may coincide with the wet season (just
ending at this time) and maximum availability of fruit such as figs.

ADDITIONAL MATERIAL. Apart from the specimens obtained by the Great
Abbai Expedition, the collections of the British Museum (Natural History) include
three further Ethiopian specimens (B.M. 28.1.11.5-7) obtained at the Donkam
River, Great Abbai, 100 miles southwest of Lake Tana, at 5,000 feet, by
R. E. Cheesman, which presumably led Ellerman, Morrison-Scott and Hayman
(1953 : 49) to include Ethiopia in the distribution of the species.

Rhinopoma hardwickei sennaarinese Fitzinger, 1866
(Plate 3 (a))

SPECIMENS, (i) Three males, three females. North eastern slope of Mount
Fantalle, Shoa. 08 58' N, 39 54' E, alt. 1,000 m. 28 September 1968.

(2) Two males, nine females. North bank of Awash River, Awash National
Park, Shoa. 08 30' N, 40 01' E, alt. 1,000 m. 25-28 September 1968.

(3) Skulls from cave deposit. Near Metahara, Awash National Park, Shoa.
08 50' N, 40 01' E, alt. 1,000 m. 28 September 1968.

TAXONOMIC NOTES. Kock (1969 : 35) has reviewed R. hardwickei with particular
reference to forms inhabiting northeastern Africa and has concluded that larger
specimens from the Sudan, Mauretania, northwest Africa, Lower Egypt, Israel,
Jordan, Aden and Sokotra should be referred to R. h. sennaariense Fitzinger, 1866,
a name thought for many years referable to R. microphyllum. Rhinopoma hard-
wickei cy stops Thomas, 1903, to which all specimens from Egypt, the Sudan and
Ethiopia were formerly referred is considered by Kock to be a smaller sub-
species inhabiting Middle Egypt and extending westwards to Hoggar and Air in the
Sahara : the small R. h. macinnesi Hayman, 1937 is thought by Kock to extend from
northern Kenya and the southern Sudan to Somalia and to Eritrea in Ethiopia.
These specimens from eastern Ethiopia are similar in size (length of forearm in
thirteen examples 52-9-56-6 (55-0)) to those which this author refers to sennaariense
from Khartoum and are considerably larger than macinnesi (length of forearm
46-6-48-4, according to Kock (pp. 45, 50)).

BATS FROM ETHIOPIA 31

COLLECTION AND FIELD NOTES. The living bats were all shot or netted inside
some small lava blister caves in the middle of the day. These provided the only
shade and cool shelter in an otherwise hot, dry and very open terrain, covered with
sparse thorn scrub, boulders and dry grass. Large numbers of Rhinopoma were
found in these caves, often hanging from the roof in small solid masses of a dozen or
more individuals. A sample of the bats was collected at random: the colonies
contained both sexes, but no signs of any young were seen. Rhinopoma was also
well represented among skeletal material removed from the floors of two of the
lava caves.

Taphozous perforatus haedinus Thomas 1915

SPECIMENS, (i) Thirteen males, eight females. "Forward Base Two", 10 km
west of Mabil, Blue Nile Gorge. 10 19' N, 36 45' E, alt. c. 1,300 m. 19 August
1968.

(2) One male, three females. North Bank of Awash River, Awash National
Park, Shoa. 08 50' N, 40 01' E, alt. 1,000 m. 25-27 September 1968.

TAXONOMIC NOTES. These specimens do not support the view (Harrison, 1961 :
150; 1962 : 763) that two closely related species of the Taphozous perforatus group
exist in Africa, one, perforatus, with dark wings, the other, Sudani, with pale or
whitish wings, the two being separated also by a small difference in the size of the
braincase, that of Sudani being on the whole very slightly larger than that of
perforatus. As noted by Kock (1969 : 74), who also rejects this view, the pigmenta-
tion of the wing membrane is variable. A number of the Ethiopian specimens have
dusky wing membranes but in others the wing membranes are translucent, especially
distally, and in this respect are closely similar to Sudani. Similarly, the dimensions
of the braincase in the Ethiopian specimens bridge the narrow interval separating
perforatus from Sudani. It is evident, therefore, that but a single species must be
recognized, a course adopted by Rosevear (1965 : 151) and by Kock (1969 : 74)
but counter to the opinion of Harrison (1961 : 150, 1962 : 763), who divided the
group into two species, perforatus having a northern and eastern distribution in
Africa, with Sudani distributed from the Sudan to southern Africa.

Subspecific classification in T. perforatus is less clear, and final clarification must,
it seems, await the advent of more material. The collections now available in the
British Museum (Natural History) suggest that a pale subspecies (T. p. perforatus
E. Geoffrey, 1818) occurring in northeastern Pakistan and in Egypt is replaced in
the Sudan and Congo by a darker, slightly larger subspecies (T. p. Sudani Thomas,
1915) which in turn in Ethiopia and Kenya gives way to an equally dark but slightly
smaller subspecies (T. p. haedinus Thomas, 1915 : according to Kock (1969 : 80)
Taphozous maritimus Heuglin, 1877 may be a prior name) extending to Somalia,
Eritrea and southwestern Arabia. Specimens from southwestern Arabia are
generally referred to T. p. haedinus but on the whole are a little paler than are
those from Ethiopia and Kenya and thus tend towards T. p. perforatus: a small
number of specimens reported as T. perforatus by Harrison (1968 : 323) from Oman

J. E. HILL & P. MORRIS

^t-

o o

r^.

Tj- OO

CO-

00^

22-

& 22-

22-

22- 22-

M
|

r>. ON

TJ- oo

CO
N

co oo

CO If

co oo
1

CO l*

oo oo

"10

^H g

o .S

nt CO

H
H

ro oo

525

Tt-

10 ^

vp~

j-.

ON ON

^ON

H-> O

!2 .S

VO 00

M-c

r T

N CO

HI
1

r r

ro ro

ON ON

g
<o

* S
o .5

HI
HI

^^ vO

H
HI

M VO

*
*
*

0)
OH
CO

'o"

'rf *m

fc.

ON ON

^ VO

>*"> CU

& &

M W

>4-l

a "~
cS

CO
CO

C^ ^N

ob oo

ro
CO

a
o

(-1

ffi

emeni

"o -S

M
HI

HI
HI

r^s vo

CO ON

,1,

N N

-j-

ON M

VO,

vO_

vO vO

^ ^2-

t* S

ho cu

op N

op op

QJ O

^- ^t-

ON N

IT) *O

iO vO

00
vO

o o

oo w

vO ^O

u~> vO

' *

"o|

H
HI

co >*

OH
CO

. 'S

perforatus
Oman*

p. haedinus
S. W. Arabi

p. haedinus
Ethiopia
p. haedinus
Kenya

p. Sudani
Sudan

p. swirae
N. Nigeria*

p. swirae
W. Africa
p. rhodesiae
Rhodesia

h h

BATS FROM ETHIOPIA 33

are said by that author to be distinctly paler than T. p. haedinus and hardly distin-
guishable in colour from material from Sudan and Cutch. Specimens from Northern
Nigeria have been separated as a distinct subspecies, T. p. swirae Harrison, 1962,
on account of their greyish coloration : Kock (1969 : 80) lists this as a synonym of
T. p. perforatus but thinks that there is a possibility that there may exist a distinct
western subspecies, for which Taphozous senegalensis Desmarest, 1820 would be the
earliest name, with swirae in synonymy. Extending in West Africa to Senegal,
Ghana, Mali and the Cameroon, T. perforatus is known also from Tanzania, Botswana
and Rhodesia, whence Harrison (1962 : 763) has described a small form,
T. p. australis (preoccupied: as a subspecies of Sudani] subsequently renamed
T. p. rhodesiae (Harrison, 19643. : 2) and considered a synonym of T. p. Sudani by
Kock (1969 : 8l). Measurements of T. perforatus appear in Table i.

COLLECTION AND FIELD NOTES. The specimens obtained near Forward Base Two
all came from some small caves which lay at the head of a stream gulley, high up the
side of the Nile Gorge. The gulley was steep, vertical in places, thickly over-grown
and passed through dense thorn scrub covering the hillsides. The caves themselves
were shallow and although dim light penetrated almost to their furthest extremities,
they provided cool shelter from the sun.

Large numbers of Taphozous were found together with single specimens of Nycteris
thebaica and Hipposideros ruber. The cave floor was thickly carpeted with guano,
and in one area, littered with the wings of large insects. These are presumed to repre-
sent the remains of insect prey caught in flight by the bats and taken back to the
roost where the thick bodies were eaten and the wings discarded. From their size and
abundance it is assumed that these insects were the prey of Taphozous, rather than
of the two smaller and less numerous bats. An endeavour was made to collect all
wings in reasonable condition from the pile and these have been identified for the
most part by Mr. Alan Brindle of the Manchester Museum, the remainder having
been examined at the British Museum (Natural History) : the species encountered
are listed in Table 2, with their relative abundance.

An attempt was made to catch all the bats in the colony and in all, 40 Taphozous
were obtained, leaving behind a few individuals that had retreated to inaccessible
parts of the cave. All the bats were sexed and weighed in the field, then 21 of the
Taphozous sample were released. The sex ratio in the full sample was 19 males:

21 females. Weights ranged from 17 gms to 23 gms, with only two animals exceeding

22 gms. The average weight was 19-5 gms with no significant sex difference.
From the narrow size range, and the presence of both sexes in equal numbers and
similar body weights it is evident that this was not a nursing colony and that
breeding must occur at some other time of year, perhaps in a different place.

The Taphozous collected in the Awash National Park were just a small sample
taken to record their presence in the lava bubble caves. Here again the cavities
were light throughout, but provided shelter and shade in an otherwise very open
habitat.

34 J- E. HILL & P. MORRIS

TABLE 2

Identities and abundance of insect wings from a roost of Taphozous perforatus
Species identified Number obtained

Orthoptera

Tettigoniidae

Diogena fausta (Burmeister) i
Mantodea

Tarachodes sp. i

Polispilota aeruginosa (Goeze) i
Lepidoptera
Saturnidae

Nudaurelea macrophthalma Ky 2

Gyanisa maja Klug i
Sphingidae

Agrius convolvuli (Linnaeus) 35

Hippotion eson (Cramer) 7

Hippotion osiris (Dalman) i

Hippotion celerio (Linnaeus) i

Nephele peneus (Cramer) i

Platysphinx stigmatica (Mabille) 2

Euchloron megaera (Linnaeus) i

Nycteris thebaica labiata (Heuglin, 1861)

SPECIMENS, (i) One male, three females. Mabil, Blue Nile Gorge. 10 20' N,
36 45' E, alt. c. 1,200 m. 18 August 1968.

(2) Two females, one male. "Forward Base Two", 10 km west of Mabil, Blue
Nile Gorge. 10 19' N, 36 45' E, alt. c. 1,000 m. 18-19 August 1968.

TAXONOMIC NOTES. These specimens are similar in size (length of forearm
41-45) to examples of N. t. thebaica from Egypt and from other localities (Gondar;
Gallabat; Ghibbey Valley, o8i5'N., 3755'E.) in western Ethiopia but are
slightly greyer ventrally. For this reason they are referred provisionally to
N. t. labiata from Keren. The subspecies of N. thebiaca are discussed by Kock
(1969 : 98).

COLLECTION AND FIELD NOTES. The Mabil specimens were shot in the roof of a
disused hut standing on a hilltop in open cultivated ground. One of the Forward
Base Two specimens was taken from the same small caves as the large sample of
Taphozous (above), the others from a similar little rock shelter lower down the
same river gulley.

ADDITIONAL SPECIMENS. The Sandhurst Ethiopian Expedition, 1966 also
obtained N. thebaica at a locality northeast of Lake Chamo, southern Ethiopia,

BATS FROM ETHIOPIA 35

where specimens were found sleeping separately in deep, shady pits in the ground.
Skulls of Nycteris sp. were also present among the cave floor bone debris collected
from certain lava blister caves in the Awash National Park in September 1968.

Cardioderma cor (Peters, 1872)

SPECIMENS. Skeletal material. Awash National Park, Shoa. o85o'N, 40
01 ' E, alt. c. 1,000 m. 28 September 1968.

COLLECTION AND FIELD NOTES. Collected among large numbers of mammal
bones found on the floor of the lava blister caves.

ADDITIONAL SPECIMEN. A skin, thought to be Cardioderma, formed part of a
small collection of local mammals held at the HQ of the Awash National Park.

Rhinolophus clivosus acrotis Heuglin, 1861

SPECIMENS. Skeletal material. Awash National Park, Shoa. o85o'N, 40
01' E, alt. c. 1,000 m. 28 September 1968.

COLLECTION AND FIELD NOTES. Subfossil cave remains, as for previous species.

Rhinolophus landeri dobsoni Thomas, 1904

SPECIMEN. One male. Temporary Base, mouth of Azir River, Blue Nile Gorge.
10 29' N, 36 25' E. alt. 1,000 m. 20 August 1968.

TAXONOMIC NOTES. This specimen agrees in size with the type specimen of
dobsoni from Kordofan and with a small series from the Sudan and western Ethiopia
in the collections of the British Museum (Natural History). The series as a whole
confirms the measurements of Kock (1969 : 175) and shows that specimens from the
Sudan and from Ethiopia are generally smaller (length of forearm in 12 examples
41-5-44-6 (42-8) than those from Malawi (R. I. lobatus: length of forearm in 16
examples 42-4-46-6 (44-8)), or from Tanzania (length of forearm in 5 examples
44-4-46-0 (45-1)). Specimens from Zambia are slightly larger than those from
Malawi (length of forearm in n examples 42-5-47-8 (45-5)) but from Kenya are
smaller (length of forearm in 10 examples 41-1-45-8 (43-9)), approaching the
northern subspecies in size.

COLLECTION AND FIELD NOTES. Caught in hand-held mist net, 20.05 hrs on
open river bank beside tall trees.

Rhinolophus hipposideros minimus Heuglin, 1861

SPECIMEN. One female. Lake Baa-sa-ka, Awash, Shoa. 08 50' N, 40 01' E,
alt. c. 1,000 m. 26 September 1968.

TAXONOMIC NOTES. This specimen appears to be the second of R. h. minimus to
be recorded from Ethiopia, the subspecies being known in northeastern Africa from
Keren (the type locality) and from Sennaar, Sudan (Andersen, 1904 : 455). The

36 J. E. HILL & P. MORRIS

specimen from Shoa agrees with the Sennaar example excepting only that it has
narrower zygomata and has the anterior upper premolar (pm 2 ) smaller, more
rounded and less angular in cross-section. Measurements (the Sennaar example
in parentheses): length of forearm 367 (357); total length of skull to canine 14-4
(14-5); condylocanine length 12-6 (127); rostral width 3-4 (3-3); zygomatic width
6'7 (7 -5); least interorbital width 1-4 (1-5); width of braincase 6-2 (6-3); mastoid
width 6-9 (7-1) ; c - m3 5-0 (5-0) ; c - m 3 5-3 (5-3).

COLLECTION AND FIELD NOTES. Caught after dark (20.00 hrs) in a hand-held
mist net as it flew low over the open, muddy lake shore.

Rhinolophus simulator Anderson, 1904

SPECIMEN. One female ; B.M. 64.854. Three miles south of Goba, Bale Province,
(presumably about 5 30' N, 40 05' E P.M.) c. 3,000 m. 1962.

TAXONOMIC NOTES. This specimen was first identified in 1964 on accession to the
collections of the British Museum (Natural History) as an example of R. hipposideros
minimus, but further examination in the course of reporting the Great Abbai
material shows that although in the features of the noseleaf it clearly resembles
this taxon, it is much too large, particularly cranially, to represent it. It is referred
to the rather larger R. simulator, not hitherto reported from any more northerly
locality than southern Tanzania, although Dr. K. Koopman (in litt.) of the American
Museum of Natural History has identified specimens from western Kenya and the
southern Sudan with this species.

The Ethiopian specimen agrees closely with simulator in the structure of the sella,
which is wide and very slightly constricted at a point a little above its centre and in
its low, rounded connecting process which rises slightly above the rounded tip of the
sella. The lateral margins of the lancet are slightly concave and the lancet itself is
rounded towards the tip to f6rm a broad point. Apart from its generally smaller
size, the skull agrees closely with that of simulator, with prominent rostral swellings,
a shallow rostral sulcus, the anterior upper premolar (pm 2 ) in the toothrow and
with the second lower premolar (pm 3 ) minute and extruded. Like simulator, it
differs from R. swinnyi and R. denti (perhaps conspecific) in larger rostral swellings,
the presence of a rostral sulcus and in having a longer supraorbital region. Rhino-
lophus alticolus Sanborn, 1936 from the Cameroon is also very like simulator, differing
chiefly in slightly larger size, rather less acuminate lancet and in having the mesop-
terygoid fossa a little wider. There can be little doubt that simulator and alticolus
are conspecific : the single specimen from Ethiopia points to the possible existence of
a generally smaller montane subspecies in that region, a matter to be resolved by
further specimens. The prior name is simulator by many years: alticolus, first
described as a subspecies of R. alcyone, has evidently no close affinity with that
species. Measurements of R. simulator are compared in Table 3.

COLLECTION AND FIELD NOTES. Collected by P. M. Barrer and presented by the
Imperial College (University of London) Ethiopian Expedition 1962.

BATS FROM ETHIOPIA

TABLE 3

Measurements of Rhinolophus simulator

M-l

-o

- i

"rt

a>
,0

I"H

^ <u
be g

<u
&

-g

G
ea

IH
QJ

tfi o
**

"3
u

H-l
O

3
'o

h-1

o -a

c
o

-->

co
O

o
bo

h-l

4->

T3
S

-i->

to
03

R. s. simulator

95-7-1-4

43-9

4'3

6-4

1 1 -6

6-9

95-7-1-5

44-2

4'5

8-7

2-O

6-7

n-5

7-0

4.12.1.4

43-8

4'3

8-8

2-2

6-6

7-0

4.12.1.5

42-7

5.12.9.89

4-8

9-1

2-2

6-9

u-8

7'3

59-355

42-5

18-8

16-6

4'5

9-1

2'3

8-9

9-4

6-7

7-0

66.5445

44'5

4'7

2-4

6-7

7-1

68.999

44-2

18-7

16-5

4-8

8-8

2-2

8-7

9-2

6-6

1 1 -8

7-0

68.1000

44'5

18-8

16-4

4'4

9-0

2-2

8-6

9-2

6-5

11-7

6-9

68.1001

42-9

18-3

16-1

4-6

8-9

2-2

8-4

9-1

6-4

"'5

6-7

68. 1 002

4 2 -9

18-5

16-4

4'5

9-0

2-2

8-3

9-0

6-5

n-3

6-9

14.6.13.2

43 '

4'3

6-6

7-0

11.4.23.1

44-2

18.0

15-9

4-4

2-2

8-4

9-0

6-2

6-5

Jf?. simulator

64.854

42-6

17-5

15-4

4'3

8-5

2-0

7'7

8-4

6-0

6-4

R. s. alticolus

56.187

46*3

18-7

16-7

4-9

2-2

9-1

6-8

7'3

56.188

46*5

18-6

16-3

4-8

8-9

2-3

8-1

9-0

6-6

7-0

68.895

45'5

n-8

7-2

68.896

45-8

19-1

16-9

4'9

9'3

2-2

8-3

9-3

6-9

1 1 -9

7-2

68.897

46-2

18-8

16-7

4'9

9'3

2< 3

8-3

9-3

6-7

11-7

7'3

68.898

44'9

18-8

4'9

8-9

2-3

6-8

11-7

7-2

Rhodesia

Transvaal
Zambia

Malawi
Tanzania

Ethiopia

Cameroon

Rhinolophus fumigatus fumigatus Riippell, 1842

SPECIMENS, (i) One female. Mouth of Fincha River, Blue Nile Gorge, 10
3' N, 37 20' E, alt. 1,000 m. 12 August 1968.

(2) One male. About 30 km southeast of "Portuguese Bridge", near Mota,
Blue Nile Gorge, 11 20' N, 38 10' E, alt. c. 1,300 m. 22 September 1968.

COLLECTION AND FIELD NOTES. Both specimens shot; the Fincha River one
coming from a small rock fissure on a rocky hillside covered with thorn scrub and
the other from a cave in a cliff beside the Nile.

38 J. E. HILL & P. MORRIS

Hipposideros caffer caffer (Sundevall, 1846)

SPECIMEN. One male. "Forward Base Two", 10 km west of Mabil, Blue Nile
Gorge, 10 19' N, 36 45' E, alt. c. 1,000 m. 18 August 1968.

TAXONOMIC NOTES. Hitherto, many authors have followed Andersen (1906 : 275)
in maintaining H. caffer as a polytypic species with a number of subspecies among
which wide intergradation occurred. However, Hollister (1918 : 85) considered
that two species, readily separable by size, were to be found together in East Africa,
and, more recently, Lawrence (1964 : i) has suggested that the relative sizes of the
narial compartments can be used as a specific distinction between these, the putative
subspecies caffer and ruber. This author did not, however, attempt to allocate the
various named forms hitherto ascribed to caffer beyond suggesting that centralis
should be associated with ruber rather than with caffer. Kock (1969 : 130, 133;
also discusses the classification of the group and concludes that two species can be
recognized. Specimens from the Great Abbai Expedition and others from Ghana
examined recently confirm the views of these authors and have prompted a further
examination of the entire complex as it is represented in the collections of the
British Museum (Natural History). The majority of specimens can be allocated
readily to one or other of two groups, as Koopman (1966 : 158) has noted. One
(caffer, angolensis, tephrus, Inanus) is composed of smaller (length of forearm usually
less than 48), generally more greyish (in the dull phase) forms with small median
posterior narial compartments and wide lateral inflations. Those allocated to the
second group (ruber, centralis, guineensis, niapu] are generally larger (length of fore-
arm usually greater than 48), browner (in the dull phase) and have larger median
posterior narial compartments with narrow lateral inflations. There is evidently
an ecological preference as is indicated by Verschuren (1957 : 354, 373 for centralis
and nanus], Lawrence (1964 : 4) and Koopman (1966 : 158), the members of the first
group occurring in the drier woodland and savannah regions, those of the second
group in wetter, densely forested areas, as suggested by Brosset (1968 : 338).
There exist, however, wide areas of sympatry on the fringes of the forest areas and
this has led to the difficulties encountered when all of the named forms are considered
to be subspecies of a single polytypic species (Hill, 1963 : 63). Members of either
group may be readily recognized over most of Africa but in northern Angola and the
Lower Congo the local representative (centralis} of the larger group is reduced in size
and distinction from that (angolensis) of the smaller group is difficult. This circum-
stance led Koopman (1966 : 158) to retain the concept of a single polytypic species.
Both species occur in two colour phases : in the dull phase caffer is greyish and ruber
brownish, while the bright phase of caffer is some shade of orange, of ruber more
rufous.

Hipposideros caffer is distributed throughout most of Africa excluding the central
forested region from Morocco to Senegal, Sierra Leone, Ghana, Senegambia, Nigeria,
northeastern Congo, Sudan, Ethiopia, Somalia, Kenya, Tanzania (including Zan-
zibar), Pemba Island, Zambia, Rhodesia, Malawi, Natal, Transvaal, Cape Province,
South West Africa, Angola and Gabon: outside Africa the species extends to the
Yemen. The following subspecies may prove valid:

BATS FROM ETHIOPIA 39

Hipposideros coffer coffer (Sundevall, 1846)

Mainly northeastern, eastern and southern Africa.

Hipposideros coffer angolensis (Seabra, 1898)

South West Africa; Angola; Gabon; Lower Congo.

Hipposideros coffer tephrus Cabrera, 1906

Northern and northwestern Africa ; drier regions of West Africa.

Hipposideros coffer nanus J. A. Allen, 1917
Northeastern Congo.

It seems that aurantiacus de Beaux, 1924 from Somalia is based on an example of
H. c. coffer in the red or brighter phase, while apparently braimo Monard, 1939 from
Portuguese Guinea should be synonymized with H. c. tephrus (Aellen, 1956 : 26;
Rosevear, 1965 : 226). Lawrence (1964 : 3) and Koopman (1965 : 10, 1966 : 158)
agree that nanus J. A. Allen, 1917 is a subspecies of H. coffer.

Specimens from the Sudan are referred to H. c. tephrus by Koopman (1965 : 10)
and Kock (1969 : 130). The Ethiopian specimen from the Great Abbai collection,
however, has a generally slightly larger skull than tephrus and consequently is
referred to H. c. coffer. Measurements; length of forearm 46-9; greatest length of
skull to canine 17-2; condylocanine length 14-9; rostral width 4-4; zygomatic
width ; least interorbital width 2-8; width of braincase 8-6; mastoid width 9-4;
c 1 -c 1 4-0; m 3 -m 3 5-8; c- m 3 5-8; length of mandible 10-1; c - m 3 6-4.

COLLECTION AND FIELD NOTES. The single specimen was caught after dark
(20.45 hrs) in a hand-held mist net as it flew low over a flat riverside sandbank in
an area of thick bush.

Hipposideros ruber centralis Andersen, 1906
(Plate 3 (b))

SPECIMENS, (i) One female. Mouth of Fincha River, Blue Nile Gorge.
10 03' N, 37 20' E, alt. 1,000 m. 12 August 1968.

(2) Two males and two females. "Forward Base Two", 10 km west of Mabil,
Blue Nile Gorge. 10 19' N, 36 45' E, alt. 1,000 m. 15-19 August 1968.

(3) One male. "Forward Base Three", mouth of Didessa River, Blue Nile
Gorge. 10 05' N, 35 38' E, alt. c. 1,000 m. 26 August 1968.

TAXONOMIC NOTES. Reasons for regarding ruber as a distinct species rather than
a subspecies of coffer are discussed above. It is of interest to note that specimens
(B.M. 67.1135-1140) collected by the Sandhurst Ethiopian Expedition, 1966 at the
southwest corner of Lake Abaya are also referable to H. r. centralis which clearly
extends some distance into Ethiopia.

4 o J. E. HILL & P. MORRIS

Hipposideros ruber is distributed through the forests and savannahs of Ethiopia,
the Sudan, Uganda, Kenya, Tanzania, Zambia, Angola, Gabon, Congo (Kinshasa),
Congo (Brazzaville), Cameroon, Fernando Poo, Nigeria, Ghana, Gambia, Sierra
Leone, Liberia, Spanish Guinea, Senegal, Sao Tome' Island and Principe Island.
Possible subspecies are:

Hipposideros ruber ruber (Noack, 1893)

Tanzania, Zambia, Angola (Sanborn, 1950 : 58).

Hipposideros ruber centralis Andersen, 1906
Ethiopia, Sudan, Uganda, Kenya.

Hipposideros ruber guineensis Andersen, 1906
West Africa.

Hipposideros ruber niapu J. A. Allen, 1917
Northeastern Congo.

Specimens reported by Aellen and Brosset (1968 : 447) as H. coffer from Congo
(Brazzaville) seem likely to represent ruber (possibly H. r. centralis) : others reported
from the Cameroon by Aellen (1952 : 72, 73) as H. c. caffer and H. c. angolensis
seem from measurements to be referable to H. ruber. Aellen (loc. cit., pp. 74, 75)
also records guineensis and ruber from the Cameroon, as subspecies of H. caffer.

COLLECTION AND FIELD NOTES. The Fincha River specimen, and three of those
from Forward Base Two were all caught after dark in a hand-held mist net as they
flew low over riverside sand banks in thick scrub habitat. The specimen from the
Didessa River was in rather less open habitat, flying around among riverside bushes
at 21.05 hrs. The remaining H. ruber from Forward Base Two was netted in the
same small caves as large numbers of Taphozous perforatus (q.v.).

Two sharply contrasted colour phases are represented in the collection
grey/brown and bright orange/red, with both forms being encountered at a single
locality (Forward Base Two).

Asellia tridens tridens (E. Geoffroy, 1818)

SPECIMEN. Skeletal material. Awash National Park, Shoa. o85o'N, 40
01' E, alt. c. 1,000 m. 28 September 1968.

COLLECTION AND FIELD NOTES. Part of a collection of mammal bones removed
from the floor of a lava blister cave.

BATS FROM ETHIOPIA 41

Asellia patrizii de Beaux, 1931

SPECIMEN. One male. North bank of Awash River, Awash National Park,
Shoa. 08 50' N, 40 01 ' E, alt. c. 1,000 m. 25 September 1968.

TAXONOMIC NOTES. This species has been known hitherto only from Danakil,
Ethiopia (the type locality) and from two other locations in Ethiopia, namely
Assab, Eritrea and Entebebir Island, near Dahlak Kebir Island, off Massawa,
Eritrea, the specimen recorded now from the Awash National Park being in fact the
first to be received at the British Museum (Natural History). It demonstrates
effectively the much smaller skull of patrizii when compared with tridens which has
been obtained (vide supra) in cave remains from the same area. Harrison (1965 : 4)
noted that specimens from Entedebir Island are slightly smaller than those
recorded from the mainland and thought therefore that they might prove to be
subspecifically separable. However, the specimen from the Awash National Park
is very similar in size to those reported by Harrison and does not support this view.
Measurements: length of forearm 40-1; greatest length of skull 14-7; condylobasal
length 13-1; condylocanine length 12-7; zygomatic width 7-6; least interorbital
width 1-9; width of braincase 6-1; mastoid width 7-1; c 1 - c 1 3-9; m 3 -m 3 5-2;

COLLECTION AND FIELD NOTES. Shot hanging from the roof of a lava blister cave
in very dry open, rocky terrain, 1500 hrs.

Triaenops persicus afer Peters, 1877
(Plate 3 (c))

SPECIMENS, (i) One male and one female. "Forward Base Three", mouth of
Didessa River, Blue Nile Gorge. 10 05 'N, 35 38' E, alt. c. 1,000 m. 28 August
1968.

(2) One female. Awash National Park, Shoa. o85o'N, 4ooi'E, alt. c.
1,000 m. 28 September 1968.

TAXONOMIC NOTES. Although recorded from Somalia and Kenya, these speci-
mens appear to be the first of Triaenops to be reported from Ethiopia. A rather
larger subspecies, T. p. majusculus, has been described by Aellen and Brosset
(1968 : 450) from the Congo (Brazzaville).

COLLECTION AND FIELD NOTES. Both specimens obtained at Forward Base
Three were caught with a hand-held mist net. The male was flying low over a
maize plot on the bank of the Nile at 19.50 hrs, the female was flying low over the
river itself at 22.00 hrs. The habitat is dense trees and bush with areas of maize
cultivation, sharply contrasting with the dry, open Awash locality where the other
specimen was shot in one of the lava blister caves in the middle of the day.

ADDITIONAL SPECIMEN. A further specimen (B.M. 69.875) collected by Mr. C.
Buer and presented by Dr. M. J. Largen has been examined recently. It was
obtained on the main road between Lake Langano and Addis Ababa.

42 J. E. HILL & P. MORRIS

Pipistrellus kuhlii fuscatus Thomas, 1901

SPECIMEN. One female. Ghimbi, Wollega. 09 10' N, 35 50' E, alt. 2,150 m.
31 September 1968.

TAXONOMIC NOTES. Mertens (1925 : 22) pointed out that africanus Ruppell,
1842 from Shoa is very similar to f^lscatus Thomas, 1901 from Kenya, for which it
is considered a prior name by Kock (1969 : 168).

COLLECTION AND FIELD NOTES. This specimen was obtained "from a house"
(presumably in the town of Ghimbi), no further information is available.

Pipistrellus nanus (Peters, 1852)

SPECIMENS, (i) One male and two females. Sabeta, Shoa. o855'N, 38
40' E, alt. 2,500 m. August 1968.

(2) One male. Ghimbi, Wollega. 09 10' N, 355o'E, alt. 2,150 m. 2 Sep-
tember 1968.

COLLECTION AND FIELD NOTES. The Sabeta specimens were collected by local
children from the axils of banana leaves. The Ghimbi animal was found by a
local boy, but no further details are available.

Eptesicus somalicus somalicus (Thomas, 1901)

SPECIMENS, (i) One male and one female. Mouth of Fincha River, Blue Nile
Gorge. 10 03' N, 37 20' E, alt. c. 1,000 m. 12 August 1968.

(2) One immature female. "Forward Base Two", 10 km west of Mabil, Blue
Nile Gorge. 10 19' N, 36 45' E, alt. c. 1,000 m. 15 August 1968.

COLLECTION AND FIELD NOTES. All specimens caught flying low over riverside
sandbanks in thick bush habitat, between 20.00 hrs and 20.30 hrs using hand-held
mist nets.

ADDITIONAL SPECIMEN. A further specimen (B.M. 67.2164) collected northeast
of Lake Abaya by the Sandhurst Ethiopian Expedition 1964 is referable to the
rather larger species E. capensis.

Glauconycteris variegata variegata (Tomes, 1861)

SPECIMEN. One female, "Forward Base Two", 10 km west of Mabil, Blue Nile
Gorge. 10 19' N, 36 45' E, alt. c. 1,000 m. 18 August 1968.

TAXONOMIC NOTES. This specimen is the first of Glauconycteris to be recorded
from Ethiopia. It has large, massive canines and cheek teeth which agree closely
with G. v. variegata rather than with the Sudanese G. v. phalaena in which the den-
tition is less massive.

BATS FROM ETHIOPIA

COLLECTION AND FIELD NOTES. By chance this specimen was found dead before
being eaten by scavengers. It was lying at the river's edge among boulders which
are subject to frequent inundation. The surrounding habitat consists of tall trees
and dense scrub. %

The following new species is described below by the senior author:

Myotis morrisi Hill, sp. nov.
(Plates i, 2 (a, b))

SPECIMEN. HOLOTYPE. B.M. 70.488. Adult female. A flat, card-mounted skin
with skull; collector's number Aio7. "Forward Base Three", mouth of Didessa
River, Blue Nile Gorge. 10 05' N, 35 38' E, alt. c. 1,000 m. 28 August 1968.

OTHER MATERIAL: none.

TAXONOMIC NOTES. DIAGNOSIS. Similar to Myotis tricolor (Temminck, 1832)
of eastern Africa but differing from this species in its generally more orange dorsal
coloration; unicolored and not bicolored ventral pelage; elongate, narrower skull
with uninflated braincase and supraorbital region, the braincase shorter, more
globular and markedly narrower than in M. tricolor, the rostrum proportionately
longer and less broadened than in that species. Dentition less massive than in
M. tricolor ; second upper premolar (pm 3 ) relatively larger and incompletely intruded
from the toothrow; second lower premolar (pms) relatively larger, not compressed
between pm 2 and pnu; posterior upper premolar (pm 4 ) with narrower lingual shelf,
separated from the lingual shelf of m 1 by a wider interspace.

DESCRIPTION. Of moderate size (length of forearm 45-4) for the genus; anterior
margin of ear smoothly convex, posterior margin concavely emarginated in its distal
half, the proximal half convex. Tragus long, its length equal to one half of the
length of the ear, tapered, with slender, rounded tip directed slightly posteriorly;
anterior margin faintly convex, especially distally, posterior margin concave distally,
convex proximally, slightly serrated, a small rounded lobe at base beneath an acute,
angular emargination just below widest point of tragus. Wing inserted at base of
first toe ; calcar strongly developed, extending along almost one half of the posterior
margin of the tail membrane ; no obvious post-calcareal lobe.

Pelage woolly, dorsally overall orange brown, individual hairs tricolored, the basal
quarter blackish brown, most of remainder creamy white, hairs tipped terminally
with bright orange brown. Dorsal pelage extending narrowly on to the wing
membrane and on to the tail membrane for nearly one half of its width. Ventral
pelage unicolorous dull creamy white, tinged faintly with brown on chin and flanks.
Wing membrane generally black but antebrachium and endopatagium pale orange
yellow, a narrow band of the same colour extending across the membrane immedi-
ately behind the forearm; anterior edge of membrane and area between first and
second metacarpals and tail membrane similarly coloured. Tibia flanked by a
narrow band of orange hairs on wing membrane and a wider band of similar hairs on
tail membrane. Toes with a sparse covering of long orange hairs.

44 J- E. HILL & P. MORRIS

Skull (Plates i, 2 (a)) elongate, the braincase not especially inflated; rostrum
narrow, the supraorbital region not expanded, the supraorbital ridge forming an
uninterrupted curve; a shallow, narrow median rostral depression. Narial emar-
gination narrow, V-shaped posteriorly, its apex rounded. Palate long, narrow,
anterior palatal emargination rounded posteriorly, extending almost to a line
joining the centres of the canines; narrow post-palatal extension; shallow basi-
occipital pits. Inner upper incisor (i 2 ) longer than wide, bicuspid, with strong
angular anterior cusp ; smaller posterior cusp extending for two thirds of the height
of the anterior cusp. Outer upper incisor (i 3 ) wider than long, bicuspid, closely
appressed to inner tooth, cusps lying transversely to line of toothrow. Outer cusp
the larger, rising from a narrow cingulum shelf to a height equal to that of the
posterior cusp of inner tooth ; supported internally by a smaller secondary cusp for
two thirds of its height, the tooth somewhat hollowed internally, separated from
the canine by a short diastema. Anterior upper premolar (pm 2 ) about as high as i 3 ,
a little larger at base, in contact with canine, a slender pointed cusp with wide,
strong cingulum. Second upper premolar (pm 3 ) rising slightly above cingulum of
pm 2 , slightly less than one half its basal area, visible externally, a simple cusp rising
from a narrow cingulum, slightly intruded from row but separating pm 2 and the
posterior upper premolar (pm 4 ), in contact with these teeth. Posterior upper
premolar (pm 4 ) with strong cusp and short, narrow lingual shelf with a wide inter-
space, separating it from the lingual shelf of the first upper molar (m 1 ). Lower
incisors imbricated, first (ii) and second (i 2 ) with four cusps, outer cusp of ii incipient,
of 12 well developed but lower than inner cusps. Lower premolars not especially
compressed: second (pm 3 ) rather more than one half the height of anterior tooth
(pm 2 ) and approximately one half its basal area, in contact with pm 2 and posterior
lower premolar (pm4), but not compressed or displaced, its length and width equal.
The measurements of the new species are compared with those of M. tricolor in
Table 4.

REMARKS. In some respects M. morrisi resembles M. bocagei but is larger, has
unicolored and not bicolored ventral pelage, a relatively longer rostrum lacking
supraorbital inflation, narrower narial and anterior palatal emarginations and
relatively narrower palate. Dentally the two species are closely similar but the
teeth of M. morrisi are more generally massive than are those of M. bocagei and
M. morrisi lacks a protoconule on the anterior ridge of the first and the second upper
molars (described in M. bocagei by Harrison (1964^ : 135).)

Tate (1941 : 539) reviewed the subgenera of Myotis and (p. 552) referred tricolor
to the subgenus Selysius. However, both tricolor and morrisi approach the sub-
genus Chrysopteron in dichromatic wing pattern, although this is less evident in
tricolor, and in the presence of four lobes on the inner (ii) and second (i 2 ) lower
incisors, but retain a relatively high braincase and concave frontal profile. I have
much pleasure in associating with this new species the name of my co-author,
Dr. Pat Morris, of Royal Holloway College, University of London, in recognition
of his many services to the study of the Ethiopian fauna while with the Great Abbai
Expedition.

BATS FROM ETHIOPIA

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SSOJ013

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46 J. E. HILL & P. MORRIS

COLLECTION AND FIELD NOTES. The specimen was caught by Messrs. Yalden,
Largen and King of the Great Abbai Expedition, using a hand-held mist net. The
bat was flying after dark over the river Nile, near its north bank, at a height of
about one metre. The surrounding habitat comprises mainly maize plots and thick
bush. The specimen was photographed whilst still alive (Plate 2 (b)).

Scotophilus sp.

SPECIMENS, (i) One subadult male. Mouth of Fincha River, Blue Nile Gorge,
10 03' N, 37 20' E, alt. 1,000 m. n August 1968.

(2) One subadult male. "Forward Base Two", 10 km west of Mabil, Blue Nile
Gorge, 10 19' N, 36 45' E, alt. c. 1,000 m. 15 August 1968.

TAXONOMIC NOTES. No attempt has been made to allocate these subadult
specimens to any of the named forms of Scotophilus in northeastern Africa.

COLLECTION AND FIELD NOTES. Both individuals were found flying over riverside
sandbanks at a height of about 2 metres, the surrounding habitat consisting mainly
of dense scrub. The Fincha River specimen was shot in flight (with a pistol!) at
19.15 hrs, the other animal was the only bat caught in a fixed, stationary mist net
(19.30 hrs) on the whole Expedition, except for bats caught at their roosts.

Miniopterus inflatus africanus Sanborn, 1936

SPECIMEN. One female. North bank of Awash River, Awash National Park,
Shoa. 08 50' N, 40 01 ' E, alt. c. 1,000 m. 23 September 1968.

COLLECTION AND FIELD NOTES. The bat was flying fairly high (6-8 metres)
among widely spaced acacia trees in open thorn scrub and grassland. It was shot
in flight at 23.00 hrs.

Otomops martiensseni martiensseni (Matschie, 1897)

SPECIMEN. One female, B.M. 69.1256. French Somaliland (Territory Afars and
Issas). 11 46' N, 42 39' E, alt. 1,471 m. 8 August 1967.

TAXONOMIC NOTES. This specimen, collected by the Sandhurst French Somali-
land Expedition, records 0. martiensseni for the first time from French Somaliland
and represents a wide extension of range from Kenya.

Tadarida pumila (Cretzschmar, 1830)

SPECIMENS, (i) One male and eight females (three of them immature) . North
bank of Awash river, Awash National Park, Shoa. o85o'N, 4ooi'E, alt. c.
1,000 m. 25-26 September 1968.

(2) Two males and four females. Ghimbi, Wollega. 09 10' N, 35 50' E, alt.
2,150 m. 1-4 September 1968.

BATS FROM ETHIOPIA

COLLECTION AND FIELD NOTES. The Awash specimens formed part of a roost of
bats, living behind the loose bark of a dead tree. A sample of the animals was
collected in the late afternoon using a mist net fixed to the tree trunk. Four of the
Ghimbi animals (two females and two males) were living in the eaves and under the
corrugated iron sheeting of the Mission Church roof, one of the others is recorded as
coming from a house and the exact origin of the sixth Ghimbi specimen is not known,
though it is likely to have been collected with the rest.

These two localities for T. pumila (Awash and Ghimbi) could not be more contrast-
ing and suggest a lack of strict habitat requirements for this bat. The Awash
locality was very hot, dry, open bush whereas Ghimbi is 1,200 m higher on the cool,
wet, heavily cultivated highland plateau. The only similarities between the two
roosts are that the bats were living close together in a narrow, cramped space and
both sexes were present in each colony.

Tadarida nigeriae nigeriae (Thomas, 1913)
(Plate 3 (d))

SPECIMEN. One male. North bank of Awash River, Awash National Park,
Shoa. 08 50' N, 40 01' E, alt. c. 1,000 m. 26 September 1968.

TAXONOMIC NOTES. There is but one previous record of T. nigeriae from Ethiopia
(Ingersol, 1968 : 60), from the Gota River in the eastern part of the country. The
species has been known hitherto from no locations nearer to Ethiopia than the
northeastern Congo and southeastern Tanzania.

COLLECTION AND FIELD NOTES. This single specimen was among a sample taken
from the roost of T. pumila mentioned above. It seems peculiar that two species
should be living behind the same small piece of tree bark in a mixed colony.

Tadarida africana (Dobson, 1876)

SPECIMEN, (i) One male B.M. 28.1.11.40. Fatam river, Great Abbai (=Blue
Nile), c. 70 km south of Lake Tana, approx. 10 25' N, 37 oo' E. alt. c. 1,900 m.
17 March 1927.

(2) One specimen B.M. 69.884. ? Vicinity of Addis Ababa.

TAXONOMIC NOTES. These specimens are the first of this large molossid to be
reported from Ethiopia, reported hitherto from no nearer locality than southwestern
Kenya.

COLLECTION AND FIELD NOTES. The Fatam River specimen was collected by
Major R. E. Cheesman during one of his survey visits to the Nile Gorge. The other
individual was found dead on a telegraph wire by employees of the Imperial High-
way Authority and presented by Dr. M. Largen of the Haile Selassie University,
Addis Ababa.

48 J. E. HILL & P. MORRIS

Tadarida acetabulosus natalensis (A. Smith, 1847)

SPECIMEN. One female B.M. 6.11.1.9. Given on the specimen label as "between
Shoa and Lake Rudolf" Southern Ethiopia.

TAXONOMIC NOTES. This single specimen represents a wide extension of range
for T. acetabulosus, known hitherto from Madagascar, Mauritius, Reunion and Natal,
the latter record by A. Smith being hitherto the only evidence of the occurrence of
the species on the African mainland. Although Thomas identified the specimen
correctly upon its arrival at the British Museum (Natural History) in 1906 the
record has remained unpublished and the specimen undisturbed in the collections
until it was noted by Mr. R. W. Hayman in 1965.

SUMMARY

The Great Abbai Expedition obtained 115 specimens of bats (including cave
remains), chiefly from the Blue Nile Gorge or from the Awash National Park. One
specimen from the Blue Nile Gorge proves to represent a new species closely allied to
Myotis tricolor, for which the name Myotis morrisi is proposed. Specimens in the
collections of the British Museum (Natural History) and also others collected by the
Expedition confirm the presence of Micropteropus pusillus in Ethiopia; Triaenops
persicus afer and Glauconycteris variegata variegata were obtained for the first time
in Ethiopia by the Great Abbai Expedition, which also obtained further specimens
of Asellia patrizii and Talarida nigeriae nigeriae; a few specimens in the British
Museum (Natural History) obtained from other sources and reported in this paper
furnish the first Ethiopian records of Rhinolophus simulator, Tadarida africana and
Tadarida acetabulosus, and of Otomops martiensseni in French Somaliland (Territory
Afars and Issas). The classification of Taphozous perforatus, Hipposideros caffer
and Hipposideros ruber is reviewed. Rhinolophus simulator Andersen, 1904 and
Rhinolophus alticolus Sanborn, 1936 are considered to be conspecific.

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BATS FROM ETHIOPIA

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Senckenbergiana 7 : 18-37.

ROSEVEAR, D. R. 1965. The bats of West Africa. London.
SANBORN, C. C. 1950. Chiroptera from Dundo, Lunda, northeastern Angola. Publicoes

cult. Co. Diam. Angola 10 : 51-62, 5 figs.
TATE, G. H. H. 1941. Review of Myotis of Eurasia. Results of the Archbold Expeditions

No. 39. Bull. Am. Mus. nat. Hist. 78 : 537-565, 2 figs.
VERSCHUREN, J. 1957. Ecologie, biologic, et systematique des Chiropteres. Explor. Pare.

Nat. Garamba, Miss, de Saeger. Inst. Pares. Nat. Congo Beige, Bruxelles, No. 7 : 1-473,

173 figs., i pi., map.

J. E. HILL,

Department of Zoology,

BRITISH MUSEUM (NATURAL HISTORY),

CROMWELL ROAD,

LONDON S.W.7.

P. MORRIS, Ph.D.,

Department of Zoology,

ROYAL HOLLOWAY COLLEGE (LTNIVERSITY OF LONDON),

ENGLEFIELD GREEN, SURREY.

PLATE i
Myotis morrisi. Skull and mandible xy-5

Bull. Br. Mus. nat. Hist. (Zool.) 21, a

PLATE i

PLATE 2

(a) Myotis morrisi. Skull and mandible X7'5

(b) Myotis morrisi.

Bull. Br. Mus. nat. Hist. (Zool.) 21,2

PLATE 2

PLATE

(a) Rhinopoma hardwickei sennaariense

(b) Hipposideros ruber centralis

(d) Tadarida nigeriae nigeriae

Bull. Br. Mus. nat. Hist. (Zool.) 21, 2

PLATE 3

(b)

(d)

Printed in England by Staples Printers Limited at their Kettering, Northants, establishment

SOME SPECIES OF PARASITIC
WORMS IN THE 'DISCOVERY'
COLLECTIONS OBTAINED IN
THE YEARS 1925-1936

S. MARKOWSKI

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 21 No. 3

LONDON: 1971

ON SOME SPECIES OF PARASITIC WORMS
IN THE 'DISCOVERY' COLLECTIONS
OBTAINED IN THE YEARS 1925-1936

STANISLAW MARKOWSKI

Pp. 51-65; 20 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 21 No. 3

LONDON: 1971

THE BULLETIN OF THE BRITISH MUSEUM

(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical series.

Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.

In 1965 a separate supplementary series of longer
papers was instituted, numbered serially for each
Department.

This paper is Vol. 21, No. 3 of the Zoological
series. The abbreviated titles of periodicals cited follow
those of the World List of Scientific Periodicals

World List abbreviation
Bull. Br. Mus. nat. Hist. (Zool.).

Trustees of the British Museum (Natural History), 1971

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued 10 September, 1971 Price 6op

ON SOME SPECIES OF PARASITIC WORMS
IN THE 'DISCOVERY' COLLECTIONS
OBTAINED IN THE YEARS 1925-1936

By S. MARKOWSKIf

MATERIAL AND METHODS

THE collection described below was made by 'Discovery' expeditions between the
years 1925 and 1936 and consists of twenty-three samples of parasitic worms taken
from the muscles and mesenteries, but mostly from the intestinal tract, of four
species of fish (Euthynnus pelamis, Coryphaena sp., Chaenocephalus aceratus and
Notothenia rossi), two species of birds (Phoebetria fusca and Chloephaga picta
leucoptera), two species of seals (Leptonychotes weddelli, Hydrurga leptonyx) and a
blue whale (Balaenoptera musculus). The material from seals was collected from
six specimens of Leptonychotes weddelli and ten of Hydrurga leptonyx.

In the material studied, twelve separate species of parasitic worms have been
recognized. Of these, two forms of Cestodes seem to represent new species. In
three cases, because of the juvenile condition of the specimens, generic determination
only was possible.

The bulk of the material was preserved in 4% formalin. Some specimens were
prepared as whole mounts, having been stained with Mayer's paracarmine. Serial
sections, cut at 8 (xm thick, were stained with Ehrlich's haematoxylin and counter
stained with erythrosin. The hosts were taken at localities in the Southern Ocean,
the precise localities are given with details of each particular species of parasite.

The author takes this opportunity in expressing his thanks to the National
Institute of Oceanography for entrusting this material to him and for providing
the necessary scientific apparatus.

Thanks are also due to Dr J. P. Harding, Keeper of the Zoology Department
of the British Museum (Natural History) for providing the writer with accommodation
and the loan of microscopical equipment, and to Mr S. Prudhoe, Mr J. W. Coles
and Mr R. A. Bray of the same Museum for their assistance in the course of this
investigation, as well as to Professor J. G. Baer, who kindly lent the original material
of Hymenolepis bisaccata Fuhrmann, 1906 for comparison.

fDr. Markowski died January 5, 1971

S. MARKOWSKI

Sies of
Nematodes

peci

O w

t/5 rpl

.s|

g-g

* 8

>SJ

c^O

"s E

^ "^

1.1

"** ^

r^ <i,

h 0?"

o g

oq-^

lashleyi Contracaecum
'rfoliatus osculatum

quadratum Contracaecum
osculatum

Is &,

Tetrabothriiis
heteroclitus

Hymenolepis
prudhoei sp. n

Diplogonopont
balaenopterae

Diphyllobothri
Glandicephalu

Diphyllobothri

O 173
cfl O
V
'o
O &

^ i

S
8

S v

^ ^

Notothen
rossi

O M
<^i vS

o ^
* s
A. <

^
^1

J -S

-S, OQ

alaenopt
usculus

PARASITIC WORMS IN THE 'DISCOVERY' COLLECTIONS
ABBREVIATIONS USED IN THE FIGURES

c cuticula

c.s. cirrus-sac

ex excretory vessel

l.m. longitudinal muscles

m.ex. median excretory vessel

o ovary

s.m. subcuticular muscles

t testis

u uterus

v vagina

v.d. vas deferens

v.g. vitelline glands

v.s. vesicula seminalis

SYSTEMATIC NOTES

The material examined contains twelve species of parasitic worms belonging to
four different groups and these are enumerated below.

I. TREMATODA

NOTOCOTYLIDAE Liihe, 1909

i. Ogmogaster antarctica Johnston, 1931

Host: Leptonychotes weddetti, intestine. Locality: Falkland Islands 15.7.1928.

Several specimens of this trematode were found in the intestine of two Weddell
seals. In one case they were attached to the walls of the intestine close to strobilae
of Glandicephalus perfoliatus.

Descriptions of this trematode have been given by Johnston (1931 and 1937).

II. CESTODA
TENTACULARIIDAE Poche, 1893

2. Tentacularia (larvae)

Host: Euthynnus pelamis: cysts in the abdominal muscles. Locality: i5io'N;
i83o'W; 15.10.1925.

Some six specimens are here recorded. Similar larval stages of a tetrarhynch
were described from the same host-species by Rennie and Reid (1912).

DIPHYLLOBOTHRIIDAE Liihe, 1910
3. Diphyllobothrium lashleyi (Leiper and Atkinson, 1914)

Host : Leptonychotes weddetti intestine. Localities : Falkland Island, and Grytviken,
South Georgia; 15-17.7.1928.

Great numbers of specimens were obtained from each of three seals.

56 S. MARKOWSKI

4. Diphyllobothrium quadratum (v. Linstow, 1892)

Host: Hydmrga leptonyx, small intestine and rectum. Localities: Grytviken,
South Sandwich Islands; South Orkney; 15.9, 18-22.1.1928 : 16.2.1931. The
material was collected from ten seal-hosts, each showing a very heavy infestation.

5. Diplogonoporus balaenopterae Loennberg, 1892

Host: Balaenoptera musculus, intestine. Locality: 6i53'S, 8732'E, 27.1.1936
('Southern Empress'). Large portions of strobila and few smaller fragments were
found in this sample.

6. Glandicephalus perfoliatus (Railliet and Henry, 1912)

Host: Leptonychotes weddelli, intestine. Localities: Falkland Islands, 15.7.1928;
Palmer Archipelago, 8.1.1935.

Parts of the duodenum of two seal-hosts were found infested with this species.
Detailed descriptions of the above-mentioned diphyllobothriid cestodes have been
given in earlier papers (Markowski, 1952 and 1955).

PTYCHOBOTHRIIDAE Liihe, 1902
7. Bothriocephalusjanickiisp.nov. Figs 1-5

Host: Coryphaena sp. ; stomach. Locality 2405'S, I546'N. 27.11.1925.

Some twenty-one fragments and eight complete worms were examined. The
strobila is about 8 cm long and very slender, about i mm broad. The scolex is
very large in relation to the rest of the body, being 5 mm long and i mm broad. It
is provided with a pair of groove-like bothridia. A neck was not observed (Fig. i).

The excretory system consists of two pairs of longitudinal vessels, two individual
canals at either side of the body. Of each pair the outward or dorsal vessel is
about 5x5 (Am in diameter and the inward or ventral vessel about
22-27 M- 111 x 14-21 (xm. Another single median longitudinal canal of about
11-13 x 5-10 [/.m is situated at the right side of the cirrus-sac and the uterine
opening. Its walls are thick and provided with cells arranged radially, as seen
in the Figs 2 and 3.

The longitudinal musculature consists of a very thin layer of fibres lying im-
mediately beneath the body-cuticula and two well-developed layers of fibres in-
serted in the parenchyma (Fig. 4). The cuticula is about 3-5 [zm thick.

The sexually-mature segment is from 0-87 mm to 0-9 mm broad, as measured
in transverse section. The genital pore, situated dorsally, leads into a shallow

PARASITIC WORMS IN THE 'DISCOVERY' COLLECTIONS

genital atrium. The cirrus-sac, measured in the same place, is 160 [Am high and
84 (Am broad, elongate, pyriform and situated dorsally (Fig. 2). There appear to
be about 75 testes, but a more precise number, it has not been possible to determine.
They are arranged in a single layer in the central part of the segment and measure
about 40-50 x 30-35 [Am (Fig. 5).

The ventrally-situated tocostoma or uterine pore leads into an atrium, which as
measured in transverse section is 68 [Am high and 40 [Am wide (Fig. 3).

The vitelline glands are disposed ventrally in a single layer between the longi-

FIG. i. Bothriocephalus janickii sp. nov. : scolex.

FIG. 2. Bothriocephalus janickii sp. nov.: cross-section of the segment showing cirrus-sac.
FIG. 3. Bothriocephalus janickii sp. nov. : cross-section of the segment showing tocostoma.

58 S. MARKOWSKI

tudinal muscles, sometimes slightly overlapping into the lateral fields of the seg-
ment. They are about 25-30 x 20 (xm (Fig. 4). The ovary is a deeply bilobed
structure situated in the middle region of the segment. The eggs are 40-42 x 28-
32 [xm.

There are five species of Bothriocephalus occurring in fish-hosts in the Southern
Hemisphere. However, the descriptions of some of them are very inadequate
(Prudhoe, 1969).

Bothriocephalus janickii sp. nov. differs from others quoted by Prudhoe (1969)
with its unusually large scolex and extremely slender strobila.

The species is named after the well-known Polish zoologist, the late Professor
C. Janicki.

HYMENOLEPIDIDAE Railliet & Henry, 1909
8. Hymenolepis prudhoei sp. nov. Figs 6-14

Host: Chloephaga picta leucoptera, rectum. Locality: Teal Inlet East Falkland,

5-3-I927-

Some sixty-five adult worms were collected. The length of the strobila in these
specimens is about 9 cm and the width 4 mm. The scolex is about 125 ^m in
length and 137 (xm in width (Fig. 6). The rostellar sac is unusually long in compari-
son with the scolex, being about 187 (jun in length and 37-5 ^m in width.

The rostellum bears a crown of eight hooks, each measuring 32 (xm. They have

FIG. 4. Bothriocephalus janickii sp. nov.: cross-section of the segment showing the

arrangement of the longitudinal muscles and vitellaria.
FIG. 5. Bothriocephalus janickii sp. nov.: horizontal section of the segment.

PARASITIC WORMS IN THE 'DISCOVERY' COLLECTIONS

a long well-developed blade, and a very thick, club-shaped handle. The guard is
weakly developed (Fig. 7). A neck was not observed. The segments are short,
elongate transversely.

The three testes, 270-212 X 112-5-125 (xm are situated posteriorly across the
segment in a single row (Figs 8 and 14). The vesicula seminalis is large and situated
in the anterior region of the segment (Fig. 9). It opens with a coiled duct into the
cirrus-sac, which measures about 130 [jan in length and 85 (j.m in width, and is
provided with thick muscular walls (Fig. 10).

The cirrus is armed with a smooth stylet, which may be observed protruding from
the genital opening.

The ovary is more or less rounded and connected with the ramifying uterus,
which in the gravid segments occupies the whole proglottis. The embryo is en-
closed in two membranes. The size of the outer membrane of the egg is 45 x 40 [j.m,
the inner membrane is 37 x 28 [zm and the embryo itself 20 x 28 (xm (Fig. 13).
Embryonic hooks were not observed.

The longitudinal muscles are well developed and form two layers, of these the
outer one is more strongly developed (Fig. 12).

I. 8

^'^{^J^^^^^^^A-^: >: J'

FIG. 6. Hymenolepis prudhoei sp. nov. : scolex.
FIG. 7. Hymenolepis prudhoei sp. nov. : hooks.
FIG. 8. Hymenolepis prudhoei sp. nov. : mature proglottis.
FIG. 9. Hymenolepis prudhoei sp. nov. : horizontal section of mature proglottis.

S. MARKOWSKI

Hymenolepis prudhoei sp. nov. which in some features may be compared with
H. Usaccata Fuhrmann, 1906, H. octacantha (Krabbe, 1869) and H. philactes
Schiller, 1951, differs in the shape and size of the hooks, which in H. bissacata are
37 (jim long, and in H. octacantha and H. philactes 32-40 (xm and 31-39 (xm
respectively. Spasski and Spasskaya (1954) give the size of the hooks in H.
octacantha: 36-38 [xm.

Although there is some similarity between the new form and the other species
mentioned, the cirrus-sac is not provided with a saccus accessorius, whilst the
stylet is smooth and the shape of the testes is also different.

Some Soviet helminthologists, namely Spasski and Spasskaya (1954), Czaplinski
(1956), Maksimova (1963), Spasskaya (1966) have erected several new genera by
breaking down the genus Hymenolepis Weinland (sensu lato).

FIG. 10. Hymenolepis prudhoei sp. nov. : horizontal section of the cirrus-sac.
FIG. ii. Hymenolepis prudhoei sp. nov.: male copulatory apparatus.

PARASITIC WORMS IN THE 'DISCOVERY' COLLECTIONS

As the erection of these new genera does not seem to have produced a clearly
understood and concise classification of the species of Hymenolepis (sensu lato),
Weinland's genus in the sense of Fuhrmann's (1932) is here accepted.

From the available literature, it seems, that the cestodes recorded from
Chloephaga picta leucoptera do not include the form described above. Avery (1966)
gives a list of parasitic worms found in this host, but this list is based entirely upon
infestation acquired under artificial conditions at Slimbridge, Gloucestershire,
where the birds are kept in captivity.

FIG. 12. Hymenolepis prudhoei sp. nov. : transverse section of the segment showing the

arrangement of longitudinal muscles.

FIG. 13. Hymenolepis prudhoei sp. nov.: egg.

FIG. 14. Hymenolepis prudhoei sp. nov. : transverse section of a segment.

S. MARKOWSKI

TETRABOTHRIIDAE Fuhrmann, 1908

9. Tetrabothrius heteroclitus (Diesing, 1850)

(Figs 15-20)

Host : Phoebetria fusca : intestine. Locality : Maiviken, West Cumberland Bay,
South Georgia 14.12.1926.

Some sixteen complete strobilae and several fragments were collected from the
intestine of a sooty albatross.

The length of the strobila is about 18 cm to 20 cm. Its anterior portion is
narrow and serrated being about 12 cm in length. The posterior part of the strobila
is 2 mm thick, coiled and shows no distinct segmentation. The mature proglottis
is 400 [Jim long and i mm broad (Fig. 16). The scolex is 290 (jun long and 330 ^m
wide, and provided with two 'auriculae' (Fig. 15). The width in that part of the
scolex is 372 (jon. The suckers are 290 [xm long and 175 [Am across. The longi-
tudinal muscles form two concentric rings. The inner one is more strongly de-
veloped (Fig. 19).

FIG. 15. Tetrabothrius heteroclitus: scolex.

FIG. 16. Tetrabothrius heteroclitus: mature segment.

FIG. 17. Tetrabothrius heteroclitus: transverse section of a mature segment.

PARASITIC WORMS IN THE 'DISCOVERY' COLLECTIONS 63

The testes, twenty-two to thirty-two, are roundish and situated in the anterior
and in the posterior part of the segment, as well as in the aporal region of the
proglottis. They are about 40 [xm in diameter (Fig. 16). The vas deferens forms
numerous coils. The cirrus-sac is 152 x 150 pm in diameter (Figs 17 and 18).

The vagina runs ventrally to the cirrus-sac. The ovary is of an irregular shape.
The egg is 45 X 37 (irn in total diameter. The diameter of the inner membrane
surrounding the embryo is 33 x 22 [jun. The embryo is 29 x 20 (j.m ; the embryonic
hooks are 12 [jun long (Fig. 20). From the available literature, it seems that
Phoebetria fusca is a new host for T. heteroditus.

FIG. 1 8. Tetrabothrius heteroditus: transverse section of a segment showing cirrus-sac.
FIG. 19. Tetrabothrius heteroditus : transverse section of a segment showing the arrangement of

the longitudinal muscles.
FIG. 20. Tetrabothrius heteroditus: egg.

64 S. MARKOWSKI

III. NEMATODA

HETEROCHEILIDAE (Railliet & Henry, 1915)
10. Contracaecum osculatum (Rud., 1802)

Host : Hydrurga leptonyx and Leptonychotes weddelli, intestine. Localities : South
Orkneys 16.2.1931; Palmer Archipelago 18.1.1935.

One male and a female specimen were found in the duodenum of H. leptonyx and
one female in the duodenum of L. weddelli.

ii. Contracaecum sp. (larvae)

Host: Chaenocephalus aceratus, liver. Locality: Sandford Bay, South Orkney,
24.1.1933.

Mass infestation of the liver of an ice-fish occurred. Because of their larval
condition the specific determination of the nematodes has not been possible.

IV. ACANTHOCEPHALA

POLYMORPHIDAE Meyer, 1931

12. Corynosoma hamanni (v. Linstow, 1892)

Host: Notothenia rossi. Locality: South Orkney 17.2.1931.
Numerous larval stages were found in the mesenteries of the host. Baylis (1929)
has given a useful description of this species.

REFERENCES

AVERY, R. A. 1966. Helminth parasites of wild fowl from Slimbridge, Gloucester. 2.

Parasites of captive Anatidae. /. Helminth 4 (3) : 269-280.
BAER, J. G. 1954. Revision taxinomique et etude biologique des Cestodes de la Famille

de Tetrabothriidae, parasites d'oiseaux de haute mer et de Mammiferes marins. Mem.

Univ. Neuchdtel 1, pp. 1-121.
BAYLIS, H. A. 1929. Parasitic Nematoda and Acanthocephala collected in 1925-1927.

'Discovery' Rep. 1 : 541-560.
CZAPLINSKI, B. 1956. Hymenolepididae Fuhrmann, 1907 (Cestoda) parasites of some

domestic and wild Anseriformes in Poland. Acta parasit. pol. 4 : 175-373, figs.
CZAPLINSKI, B. & RIZHIKOV, K. M. 1964. New data on Parabisaccanthes philactes (Schiller

1951) Spassky et Reznik, 1963 (Cestoda, Hymenolepididae) from Poland and the Lena

Delta. Acta parasit. pol. 12 : 363-371.
DUBININA, M. N. 1953. Cestodes of birds nesting in Western Siberia. Parazit. Sb. 15 : 117-

233-

FUHRMANN, O. 1906. Die Hymenolepis-Arten der Vogel. Zentbl. Bakt. ParasitKde (Orig).

41 : 352-358, 440-452, 39 figs.
1932. Les Tenias des oiseaux. Mem. Univ. Neuchdtel 8 : 381 pp.

PARASITIC WORMS IN THE 'DISCOVERY' COLLECTIONS 65

JOHNSTON, T. H. 1931. New trematodes from the Subantarctic and Antarctic. Aust, J.
exp. Biol. med. Sci. 8 : 91-98.

1937. Trematoda. Scient. Rep. Australian, antarct. Exped. 10 : 1-29.

JOHRI, G. N. 1960. Studies on some Cestodes parasites. IV. On four new species including

a new genus belonging to the Family Hymenolepididae. Proc. natn. A cad. Sci. India B

30 : 192-202.

JOYEUX, CH. & BAER, J. G. 1936. Cestodes. Faune Fr. 30 : 1-613.
LAP AGE, G. 1961. A list of the parasitic Protozoa, Helminths and Arthropods recorded

from species of the Family Anatidae (Ducks, Geese and Swans). Parasitology 51 : 1-109.
MCDONALD, M. E. 1969. Annotated bibliography of Helminths of Waterfowl (Anatidae).

Spec, scient. Rep. U.S. Fish Wildl. Serv. 125 : 1-333.
MAKSIMOVA, P. 1963. New species of Cestodes occurring in Kazakhstan swan. Trud.

Inst. Zool. Alma-Ata 19 : 126-132.
MARKOWSKI, S. 1952. The Cestodes of seals from the Antarctic. Bull. BY. Mus. not. Hist.

(Zool.) 1 : 125-150, 21 pis.

J 955- Cestodes of whales and dolphins from the 'Discovery' collections. 'Discovery'

Rep. 27 : 377-395-

NERADOVA, J. 1966. Contribution to the knowledge of the Helminthofauna of domestic

ducks (Anas platyrhyncha domestica L.) in the western parts of Czechoslovakia. VSst.

6sl. Spol. Zool. 30 : 247-255.
PRUDHOE, S. 1969. Cestodes from fish, birds and whales. Rep. B.A.N.Z. antarct. Res.

Exped. (B) 8 : 171-193-
RAUSCH, L. R. & FAY, H. F. 1966. Studies on the Helminth fauna of Alaska. XLIV.

Revision of Ogmogaster jdgerskiold, 1891 with a description of O. pentalineatus sp. n.

(Trematoda: Notocotylidae) . /. Parasit. 52 : i, 26-38.
RENNIE, S. & REID, D. 1912. The Cestoda of the Scottish National Antarctic Expedition.

Rep. Scient. Results Scott, natn. antarct. Exped. Zool. 6 : 243-255, pis.
SCHILLER, E. L. 1951. Studies on the Helminth fauna of Alaska. VI. The parasites of

Emperor Goose (Philactes canagica L.) with the description of Hymenolepis philactes n.

spec. /. Parasit. 37 : 217-229.

1952. Studies on the Helminth fauna of Alaska. III. Hymenolepis kenaiensis n.

spec. A Cestode from the Greater Scaup (Ay thy a marila nearctica) with the remarks on
endemicity. Trans. Am. microsc. Soc. 71 : 146-149.

SKRYABIN, K. I. & MATEVOSYAN, E. M. 1945. Hymenolepidid Cestodes of domestic and game

fowl of economic importance. Moscow. 488 pp., Illust.
SPASSKAYA, L. P. 1966. Cestoda of birds U.S.S.R. Hymenolepididae. Akad. Nauk. Moldav.

S.S.R. Inst. Zool., 698 pp.
SPASSKI, A. A. 1963. Hymenolepididae-tapeworms of wild and domestic birds. Part I.

Principles of Cestodology 2. Akad Nauk S.S.S.R. Moscow, 418 pp.
SPASSKI, A. A. & SPASSKAYA, L. P. 1954. Construction of the Hymenolepidids system.

Parasites of birds. Trudy gel'mint. Lab. 7 : 55-119, 27 figs.
STEFANSKI, W. 1933. Constantin Janicki un distingue Parasitologue Polonais (1876-1932).

Folia morph. 4 : 220-229, phot.
SZPOTANSKA, I. 1931. Quelques especes nouvelles ou peu, connues des Hymenolepididae

Fuhrmann (Cestodes). Pr. zool. pol. panst. Muz. przyr. 9 : 247-266.

Dr. S. MARKOWSKI

c/o Department of Zoology

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MITES\<tF THE GENUS HYPOASPIS

CANESTRINI, 1884 S.STR.

AND RELATED FORMS

(ACARI: MESOSTIGMATA)

ASSOCIATED WITH BEETLES

M. COSTA

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 21 No. 4

LONDON : 1971

MITES OF THE GENUS HYPOASPIS
CANESTRINI, 1884 S.STR. AND RELATED FORMS

(ACARI: MESOSTIGMATA)
ASSOCIATED WITH BEETLES

MICHAEL COSTA

Kibbutz Mishmar Haemek, Israel

Pp. 67-98; 101 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 21 No. 4

LONDON: 1971

THE BULLETIN OF THE BRITISH MUSEUM

(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical series.

Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.

In 1965 a separate supplementary series of longer
papers was instituted, numbered serially for each
Department.

This paper is Vol. 21, No. 4 of the Zoological
series. The abbreviated titles of periodicals cited follow
those of the World List of Scientific Periodicals.

World List abbreviation
Bull. Br. Mus. not. Hist. (Zool.).

Trustees of the British Museum (Natural History), 1971

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued 10 September 1971 Price 1-10.

MITES OF THE GENUS HYPOASPIS
CANESTRINI, 1884 S.STR. AND RELATED FORMS

(ACARI : MESOSTIGMATA)
ASSOCIATED WITH BEETLES

By MICHAEL COSTA

INTRODUCTION

RECENTLY Dr C. Athias-Henriot (Laboratoire de Faune du Sol de 1'I.N.R.A.,
Dijon) forwarded to me mite material collected from the scarabaeid beetles Oryctes
rhinoceros L. and Oryctes monoceros Ol. The beetles and mites were laboratory
reared and collected by Dr M. J. Stelzer and Mr B. Zelazny (both from U.N./S.P.C.
Rhinoceros Beetle Project, Apia, Western Samoa). Mr Zelazny informed me that
the mites feed on the beetles' eggs and may therefore have a possible role in the
biological control of Oryctes rhinoceros, a main pest of Coco-nut palms. According
to Mr Zelazny, the mites originally associated with 0. monoceros, imported from the
Ivory Coast, were even more avid egg feeders on the eggs of 0. rhinoceros than
the locally collected mites. One batch of mites, ex 0. rhinoceros, was tentatively
determined as Coleolaelaps rhinocerotis (Ouds.), although I was puzzled by the
report on their feeding behaviour. It was generally assumed that mites of the genus
Coleolaelaps Berlese, 1914 are harmless exudate feeders (Grandi, 1925; Vitzthum,
1940-43). A closer examination of material collected by myself from phytophagous
scarabaeids in Israel, material collected by Dr M. Remillet (O.R.S.T.O.M., Centre
d'Adiopodoume, Abidjan, Ivory Coast) and material from melolonthine bettles
in the U.S.A., showed that 'Coleolaelaps' served actually as a 'dumping ground'
for a number of different genera of mites associated with beetles. The mites of
the genus Coleolaelaps Berlese, 1914 have been dealt with in a separate study (Costa
& Hunter, in press) which has shown that Coleolaelaps is not closely related to
Hypoaspis Can. The present paper will deal with the genus Hypoaspis s. str.
and some additional forms.

The confusion between Coleolaelaps s. str. and Hypoaspis s. str. seems to have
arisen from the fact that both have long 'wavy' setae on the idiosoma as well as
macrosetaeon the legs, and both are associated with phytophagous lamellicorn beetles.
I should like to point out that the 'wavyness' mentioned by many authors is an
artefact of the preparation, the materials used causing apparently a slight contrac-
tion of the setal core. In living or alcohol-stored specimens the setae are straight
or slightly curved.

Mites of the Hypoaspidinae are generally considered to be the most primitive
group in the Laelapidae (Vitzthum, 1940-43; Evans, 1958) but the taxonomic
treatment of Hypoaspis Can. s. lat. remains controversial. This has been shortly
discussed by Hunter & Costa (in press), who retain at full generic status many of

7 o M. COSTA

the subgenera of Hypoaspis which have been recorded, though not used, by Evans
& Till (1966). The present study supports this view and proposes to show that
mites of the genus Hypoaspis s. str. are well defined morphologically as well as
ecologically in their host associations. A close examination of the symbiontic
mites showed a high degree of host specificity and probably in the past several
species have been confused with either Hypoaspis krameri Can. (the type species
of the genus) or Hypoaspis integer Berlese sensu Samsinak, 1960, both of which have
been also confused with each other.

Evans & Till (op. cit.) have recently described and figured both sexes of Hypoaspis
krameri from specimens associated with Lucanus sp. in Great Britain. Their
description agrees with the details which can be learned from the descriptions of
H. krameri by G. & R. Canestrini (1881), G. Canestrini (1885) and Berlese (1892),
making the two undoubtedly congeneric. The host association of the British material
makes it debatable if this is actually conspecific with the original H. krameri which
is associated with Oryctes nasicornis L. (compare discussion).

In view of the present study a redefining of Hypoaspis s. str. seemed to be
necessary.

DEPOSITION OF TYPES

The holotypes are deposited in the British Museum (Nat. Hist.). Paratypes
will be deposited in The American Museum of Natural History; The Acarina col-
lection, Department of Entomology, University of Georgia and the author's col-
lection.

Hypoaspis Canestrini s. str.

Hypoaspis Canestrini, 1884, Atti R. 1st. veneto Sci. (6) 2 : 1569; 1885, Acarofauna Ital. part I : 55.
TYPE: Gamasus krameri Canestrini, 1881.

FEMALE : Dorsal shield entire, oval, with basically 37 pairs of setae (20 podonotal
and 17 opisthonotal, fig. i). Setae i2, 54-6 considerably longer than remaining
setae, setae Z4 extremely long, usually longest idiosomal setae. A tendency exists
towards diminishing the number of setae, e.g. in Hypoaspis integer Berlese setae 73
are absent in most specimens; in Hypoaspis phyllognathi sp. n. seta 53 may be absent
on one or both sides and Hypoaspis remilleti sp. n. lacks setae 33 in all specimens.
Gnathosoma with six rows of deutosternal denticles, setae Hyp. 3 very long, distinctly
longer than remaining gnathosomal setae (fig. 7). Sternal shield hexagonal with
distinct anterior border. Genital shield tongue-shaped with marginally inserted
genital setae. Paranal setae always longer than postanal seta. Peritreme extends
anteriorly beyond the margin of coxa I, not attached to dorsal shield. Tarsus II
with two subterminal stout, blunt, spur-like setae (all and pl\, fig. 4), leg IV with
macrosetae on femur (adi), genu (ad\ t in some species this seta might be similar
in length to the remaining setae of the segment) and tarsus (adz, Pdz and pdz, fig. 6).
Macrosetae are also present on femur II (pdi) and femur III (ad\). Leg chaetotaxy
as recorded for free-living laelapids (Evans, 1963).

HYPOASPIS MITES AND BEETLES 71

MALE: With long slender spermadactyl, curved distally (fig. 14). Holoventral
shield with 10 pairs of setae in addition to anal setae (various degrees of erosion
may separate the anal shield completely in Hypoaspis integer). Peritreme anteriorly
fused with dorsal shield. Leg II with ventral stout, pointed, spine-like setae on
femur to tarsus (fig. 38). Remaining characters as in female.

The mites are usually associated with phytophagous scarabaeids, mainly Dyna-
stinae.

Hypoaspis neokrameri sp. n.

FEMALE : Dorsum covered by single dorsal shield (735 (xm long and 445 |xm wide)
with 37 pairs of setae. Podonotal setae iz, si, 54-6 are elongate and distinctly
longer than the remaining podonotal setae (fig. i). The longest dorsal setae are
Z4 (220 (xm), setae J5 straight and short (30 (xm). The shield is nearly devoid
of ornamentation. Tectum (fig. 3) triangular with denticulate proximal margins.

Tritosternum normal with pilose laciniae. Sternal shield (170 |xm long and
150 (Am wide at St2) well ornamented, posterior margin slightly convex and irregular
(fig. 2). Sternal setae long, reaching to or beyond the bases of consecutive setae.
Genital shield (distance between genital setae 105 [xm) ornamented and tongue-
shaped, metapodal shields narrow. Paranal setae distinctly longer than postanal
seta. Peritreme extending anteriorly slightly beyond the middle of coxa I, free
anteriorly and posteriorly.

Gnathosoma (fig. 7) with well-sclerotized corniculi and fimbriate internal malae.
Six rows of deutosternal denticles (8-14 per row). Movable digit of chelicera
(fig. 5) bidentate, fixed digit with one stout tooth and about ten small denticles.

The approximate lengths of the legs (excluding pretarsi) are: I 660 (xm; II
540 (Jim ; III 600 (xm ; IV 850 (xm. Tarsus II (fig. 4) with two blunt distal spines,
leg IV with macrosetae on the femur (220 (xm), genu and tarsus (fig. 6). Macrosetae
present also on femur II (150 (xm) and III (no (xm). Leg chaetotaxy normal.

MALE: Unknown.

DIFFERENTIAL DIAGNOSIS : H. neokrameri sp. n. can be separated from other
species of the complex by its long sternal shield which is longer than wide. Associ-
ated with Oryctes nasicornis L. (Scarabaeidae : Dynastinae).

MATERIAL: Holotype: , ex Oryctes nasicornis L., Tivon, Israel, May 25, 1965,
coll. M. Costa. Paratypes: 5$?, ibid.

Hypoaspis pentodoni sp. n.

FEMALE: Dorsum covered by a single dorsal shield (680 (xm long and 390 |xm
wide) with 37 pairs of setae. Podonotal setae 12, 54-6 are elongate and distinctly
longer than the remaining podonotal setae (fig. 8). The longest dorsal setae are
Z4 (220 (xm). The shield is nearly devoid of ornamentation, sites of muscle attach-
ments are striated. Tectum (fig. 12) triangular with denticulate margins.

M. COSTA

FIGS 1-7. Hypoaspis neokrameri sp. n., female. Fig. i. Dorsum. Fig. 2. Venter.
Fig. 3. Tectum. Fig. 4. Tarsus II. Fig. 5. Chelicera. Fig. 6. Leg IV. Fig. 7.
Gnathosoma, ventral view.

HYPOASPIS MITES AND BEETLES

Tritosternum normal with pilose laciniae. Sternal shield wider than long (125 [xm
long and 150 ^m wide at St2), its posterior margin is nearly straight with two small
but characteristic projections (fig. 9). Sternal setae long, extending beyond the

FIGS 8-15. Hypoaspis pentodoni sp. n., female. Fig. 8. Dorsum. Fig. 9. Venter.
Fig. 10. Leg IV. Fig. n. Tarsus II. Fig. 12. Tectum. Fig. 13. Chelicera.
Fig. 15. Gnathosoma, ventral view. Male. Fig. 14. Chelicera.

74 M. COSTA

bases of consecutive setae. The ornamented genital shield is tongue shaped (dis-
tance between genital setae no (xm). Metapodal shields narrow. Postanal seta
distinctly shorter than paranal setae. The peritreme extends anteriorly beyond the
middle of coxa I.

Gnathosoma (fig. 15) with well-sclerotized corniculi and fimbriate internal malae,
six rows of tiny deutosternal denticles. Movable digit of chelicera (fig. 13) bidentate,
fixed digit with small denticles.

The approximate lengths of the legs (excluding pretarsi) are: I 650 (xm; II
540 (xm; III 530 fxm; IV 750 [xm. Tarsus II (fig. n) with two blunt distal
spines. Leg IV (fig. 10) with macrosetae on femur (210 (Am), genu and tarsus
(^3165 (Jim).

MALE : The single male specimen is smaller than the female (dorsal shield 580 (j.m
long and 330 [xm wide). Venter covered by well-ornamented holo ventral shield
with 10 pairs of setae in addition to the anal setae. Chelicera (fig. 14) with slender,
distally curved, spermadactyl. Remaining characteristics as in female, except
peritreme which is attached anteriorly to the dorsal shield.

DIFFERENTIAL DIAGNOSIS: This species can be separated from H. neokrameri sp. n.
by its short sternal shield and from H. phyllognathi sp. n. by its shorter dorsal
setae. Associated with Pentodon bispinosus Kiist (Scarabaeidae : Dynastinae).

MATERIAL: Holotype: 9- ex Pentodon bispinosus Kiist, Mishmar Haemek, Israel,
Sept. 21, 1962 coll. M. Costa. Paratypes: all from the same host and locality:
i<$, i$, May 19, 1965; 5$$, Sept. 21, 1962; 6.?, Sept. 12, 1966; i$, Nov. 8, 1965.

Hypoaspis phyllognathi sp. n.

FEMALE : Dorsum covered by single dorsal shield (840 {xm long and 470 [xm wide)
which is ornamented with small polygons mainly in the posterior area. Regularly
37 pairs of simple setae are inserted on the shield, but in several specimens setae
53 are absent on one or both sides. Podonotal setae i2 and 54-6 are distinctly
longer than the remaining podonotal setae. The longest dorsal setae are Z4
(320 (xm), setae J5 straight and short (30 fxm). The distribution and relative
lengths of the setae are shown in fig. 16. Tectum (fig. 21) triangular, proximal
margins deeply denticulate.

Tritosternum normal with pilose laciniae. Sternal shield (145 (xm long and
155 [xm wide at St2) well sclerotized, only faintly ornamented. Sternal setae
very long, St2 extends beyond the posterior margin of the shield (fig. 17). The
tongue-shaped genital shield (distance between genital setae (125 (xm) has nearly
parallel sides and is well ornamented. The anal shield is rounded anteriorly, the
postanal seta is distinctly shorter than the paranal setae. The anal shield is flanked
posteriorly by a pair of long setae. Metapodal shields small, irregular in shape.
The peritreme extends anteriorly to the anterior margin of coxa I, it is free both an-
teriorly and posteriorly.

Gnathosoma (fig. 18) with well-sclerotized corniculi and fimbriate internal malae,
with six rows of minute deutosternal denticles. Movable digit of chelicera (fig. 22)

HYPOASPIS MITES AND BEETLES

FIGS 16-22. Hypoaspis phyllognathisp.n., female. Fig. 16. Dorsum. Fig. 17. Venter.
Fig. 18. Gnathosoma, ventral view. Fig. 19. Tarsus II. Fig. 20. Leg IV. Fig. 21.
Tectum. Fig. 22. Chelicera.

76 M. COSTA

bidentate, fixed digit with 8 small denticles in addition to a large tooth which is
associated with the pilus dentilis.

The approximate lengths of the legs (excluding pretarsi) are: I 730 [xm; II
570 (xm; III 600 (Am; IV 870 jjun. Tarsus II (fig. 19) with two dorsal distal
blunt spines, leg IV (fig. 20) with macrosetae on the femur (300 |xm), genu (210 [xm)
and tarsus. Macrosetae are also present on femora II and III, leg chaetotaxy
normal.

MALE: Unknown.

DIFFERENTIAL DIAGNOSIS: This species can be recognized by its long dorsal
setae, its large size and by the long seta adi (210 [xm) on genu IV. Associated
with Phyllognathus silenus F. (Scarabaeidae : Dynastinae) .

MATERIAL: Holotype: $, ex Phyllognathus silenus F., Bardawil, Northern Sinai,
April 10, 1968, beetle coll. H. Sandier. Paratypes: 13 <j><j>, same data; 6 $<j>, Carmia,
Israel, Nov. 16, 1966; I -, Tivon, Israel, May 6, 1966; 2 .., Ein Yahav, Israel,
May i, 1968.

Hypoaspis integer Berlese, 1911 sensu Samsinak, 1960

Laelaps (Hypoaspis} integer Berlese, 1911, Redia 7 : 186.

? 'Coleolaelaps integer Willmann, 1935, Bull. Mus. R. Hist. nat. Belg. n : 23-25, figs 14-6.
Coleolaelaps integer (male) Samsinak, 1960, Cas. dsl. Spol. ent. 57 (3) : 280-82, figs 1-6.

FEMALE : Dorsum covered by single dorsal shield (820 pan long and 430 [xm wide)
with 36 37 pairs of setae. Of 29 investigated specimens, setae Z3 were completely
absent in 19 specimens, unilaterally present in 8 and bilaterally present in 2 speci-
mens only. Podonotal setae 12, si, 54-6 are elongate, the longest dorsal setae are
Z4 (280 |xm) and a pair of postero-marginal setae (200 |xm) which are inserted on
the soft integument. The shield is devoid of distinct ornamentation, the distri-
bution and relative lengths -of the setae are shown in fig. 23. Tectum (fig. 25)
triangular with denticulate margin.

Tritosternum normal with pilose laciniae. Sternal shield (150 [xm long and
150 [xm wide at St2) ornamented with short sternal setae which do not reach the
bases of the consecutive setae (fig. 24). Metasternal setae inserted on integument.
The large genital shield (distance between genital setae 130 jxm) expands slightly
beyond the genital setae and is ornamented. Metapodal shields narrow, kidney-
shaped. Anal shield with semicircular anterior border, postanal seta distinctly
shorter than paranal setae. The anal shield is flanked posteriorly by a pair of long
(300 (xm) setae. The narrow peritreme extends anteriorly to the middle of coxa I.

Gnathosoma (fig. 29) with well-sclerotized corniculi and fimbriate internal malae,
six rows of tiny deutosternal denticles (about 12 per row). Movable digit of chelicera
(fig. 27) bidentate, fixed digit with one large tooth and about 10 sub-equal small
denticles.

The approximate lengths of the legs (excluding pretarsi) are: I 680 |xm; II
620 (xm; III 580 (xm; IV 960 [xm. Tarsus II (fig. 28) with two dorsal distal
blunt spines, with ventral setae markedly stouter than the dorsal setae. Leg IV

HYPOASPIS MITES AND BEETLES

(fig. 26) with macrosetae on femur (360 ^m), on the genu (ad-\_ 250 (xm, pdi
180 (jon) and tarsus; macrosetae are also present on femur II (270 fxm) and III
(260 (Am) . Leg chaetotaxy normal for the genus.

FIGS 23-29. Hypoaspis integer Berlese, female. Fig. 23. Dorsum. Fig. 24. Venter.
Fig. 25. Tectum. Fig. 26. Leg IV. Fig. 27. Chelicera. Fig. 28. Tarsus II.
Fig. 29. Gnathosoma, ventral view.

78 M. COSTA

MALE: The male has been described by Samsinak (1960). The ventral sclerotiza-
tion is extremely variable, out of 14 male specimens only one had a complete holo-
ventral shield, in one specimen this was eroded but the anal shield was still broadly
connected to the ventral shield and in 12 specimens, with variously shaped genito-
ventral shields, the anal shield was completely separate. In the males the cor-
responding macrosetae are longer than in the females.

DIFFERENTIAL DIAGNOSIS: The long macrosetae on femora II and III, as well as
the two macrosetae on genu IV are good diagnostic characters for this species.
Associated with Oryctes nasicornis L. (Scarabaeidae : Dynastinae) and Polyphylla
fullo L. (Scarab.: Melolonthinae).

NOTES: The original association seems to be with 0. nasicornis, the association
with P. fullo appears to be secondary (vide Costa & Hunter, in press). The speci-
mens ex P. fullo (the beetles were kindly loaned by the American Museum of Natural
History) were collected from beneath the elytra by methods described by Costa
& Hunter (op. cit.).

MATERIAL: 2 $$, 28 $$, ex Oryctes nasicornis, Bohemia, Liblice, July 16, 1960,
coll. and det. K. Samsinak; 5 <$J, 53 ?$ Polyphylla fullo, S. Russia; 3 <$<$, 41 $?,
ibid., Prussia; 4c&? 56 $$, ibid., Germany, 1897.

Hypoaspis rhinocerotis Oudemans, 1925

Hypoaspis rhinocerotis Oudemans, 1925, Ent. Ber., Amsl. 7 (146) : 30.

Coleolaelaps rhinocerotis Oudemans, 1927, Zoo/. Meded. Leiden 10 (4) : 189-193, figs 8-15.

FEMALE: Dorsum covered by a single dorsal shield (770-830 (im long and 470-
540 (Jim wide), with 37 pairs of simple setae. Setae si subequal in length with vertical
setae (ii) ; the longest podonotal setae being i2, 54-6 (35 being shorter than either
54 or s6). The central dorsal setae, especially on the opisthonotum are short and
do not extend to the bases of the consecutive setae. Setae Z4 are the longest
dorsal setae (300 (Am), setae J5 are short (30 [xm). The distribution and the relative
lengths of the setae are shown in fig. 30. Tectum (fig. 32) triangular with denticulate
margin.

Tritosternum normal with well developed laciniae. Sternal shield (170 fun
long and 185 (Jim wide at St2) only slightly ornamented. Sti distinctly shorter
than St2~4 which are long and extend markedly beyond the bases of the consecutive
setae (fig. 31). The posterior margin of the shield is irregular. Genital shield
(distance between genital setae 130 (i,m) tongue-shaped and nearly devoid of orna-
mentation. Metapodal shields narrow and irregular. Anal shield small, postanal
seta distinctly shorter than paranal setae. Peritreme extends anteriorly slightly
beyond the middle of coxa I, it is free anteriorly and posteriorly.

Gnathosoma (fig. 36) with well-sclerotized corniculi and fimbriate internal malae,
six rows of deutosternal denticles (12-18 per row). Chelicera (fig. 35) with bidentate
movable digit with a large distance between the two teeth, fixed digit slightly curved
and sickle shaped with one stout tooth and about 12 small denticles.

HYPOASPIS MITES AND BEETLES

The approximate lengths of the legs (excluding pretarsi) are: I 750 [xm; II
660 (Am; III 700 [xm; IV 950 (j.m. Tarsus II (fig. 34) with two dorsal blunt
spines, its ventral setae being much stouter than its dorsal setae. Leg IV (fig. 33)
with macrosetae on femur (270 fxm) and tarsus only, ad\ of the genu being only
slightly longer than the remaining setae on the segment. Macrosetae are present
also on femora II and III. Leg chaetotaxy normal.

FIGS 30-36. Hypoaspis rhinocerotis Oudemans, female. Fig. 30. Dorsal shield. Fig. 31.
Venter. Fig. 32. Tectum. Fig. 33. Leg IV. Fig. 34. Tarsus II. Fig. 35. Cheli-
cera. Fig. 36. Gnathosoma, ventral view.

M. COSTA

MALE: Dorsal shield smaller (740 [j,m long and 480 (im wide) than in female,
with the same chaetotaxy. The venter (fig. 37) is covered by a well- ornamented
holoventral shield with 10 pairs of setae in addition to the regular anal setae.
Remaining ventral features as in female.

500,

FIGS 37-43. Hypoaspis rhinocerotis Oudemans. Male, Fig. 37. Venter. Fig 38. Leg
II. Fig. 40. Chelicera. Deutonymph. Fig. 39. Dorsal shield. Fig. 43. Venter.
Protonymph. Fig. 41. Dorsum. Fig. 42. Venter.

HYPOASPIS MITES AND BEETLES 81

Movable digit of chelicera (fig. 40) monodentate with long slender spermadactyl
which is distally curved. Fixed digit with about 4 teeth. Leg chaetotaxy as in
female, leg II (fig. 38) with several stout pointed spine-like setae on femur to tarsus,
similar to the condition found in H. krameri (as figured by Evans & Till, 1966).

DEUTONYMPH : Dorsal shield (670 [xm long and 380 [xm wide) deeply incised later-
ally, chaetotaxy and other features as in female although several central setae
are longer (fig. 39). Sternal shield (fig. 43) extends only slightly beyond the posterior
margin of coxa IV. A number of small platelets are present on the integument.
Remaining characters as in female.

PROTONYMPH : The idiosoma (660 (xm long) is covered by two dorsal shields and
three pairs of platelets in the mesonotal region. The podonotal shield (365 |xm
long and 325 [xm wide) with n pairs of long setae all of which extend beyond the
bases of consecutive setae. The opisthonotal shield with 8 pairs of setae, 85,
Z4~5 and J5 having the same relative lengths as in the adult. The distribution and
relative lengths of the setae are shown in fig. 41. The venter (fig. 42) with a sternal
shield with 3 pairs of setae. Peritremes rudimentary.

DIFFERENTIAL DIAGNOSIS: The short central dorsal setae and the large sickle-
shaped chelicera separate H. rhinocerotis from the preceding species. Associated
with Oryctes rhinoceros L. (Scarabaeidae : Dynastinae).

MATERIAL: Numerous specimens, ex Oryctes rhinoceros, Apia, W. Samoa, July,
1969; additional specimens from eggs of 0. rhinoceros and laboratory cultures.
All the specimens were made available through the courtesy of Mr B. Zelazny and
Dr M. J. Stelzer.

Hypoaspis athiasae sp. n.

FEMALE: Dorsal shield (770-820 [xm long and 490-510 fxm wide) covers most of
the dorsum, with 37 pairs of setae. The shield is slightly ornamented, mainly on
its posterior part. Setae i2, 54-6 are the longest podonotal setae, 55 being markedly
shorter than either 54 or s6. Setae Z4 are the longest dorsal setae (280 |xm), J5
are short (30 [xm). The distribution and the relative lengths of the setae are shown
in fig. 44. Tectum (fig. 46) triangular with denticulate margin.

Tritosternum with well-developed laciniae. Sternal shield (195 [xm long and
175 [Jim wide at St2) without apparent ornamentation, with long sternal setae
which extend beyond the base of the consecutive setae. Posterior margin of shield
nearly straight and slightly irregular. Genital shield (distance between genital
setae 120 (xm) tongue-shaped and well ornamented. Narrow, kidney-shaped meta-
podal shields. The wide peritreme extends anteriorly to the anterior margins of
coxa I. The postanal seta is shorter than the paranal setae.

Gnathosoma (fig. 50) with well-sclerotized corniculi and fimbriate internal malae,
with six rows of deutosternal denticles (10-14 per row). Movable digit of chelicera
(fig. 48) bidentate, fixed digit with about 14 small denticles in addition to a larger
tooth which is associated with the pilus dentilis.

M. COSTA

The approximate lengths of the legs (excluding pretarsi) are: I 840 |xm; II
675 fxm; III 710 (xm; IV 800 {xm. Tarsus II (fig. 47) with two dorsal distal
blunt spines. Leg IV (fig. 49) with macrosetae on femur (280 fxm) and tarsus

FIGS 44-50. Hypoaspis athiasae sp. n., female. Fig. 44. Dorsal shield. Fig. 45.
Venter. Fig. 46. Tectum. Fig. 47. Tarsus II. Fig. 48. Chelicera. Fig. 49. Leg
IV. Fig. 50. Gnathosoma, ventral view.

HYPOASPIS MITES AND BEETLES

only, adi of the germ being only slightly longer than the remaining setae of the
segment. Macrosetae also present on femora II and III.

MALE: The dorsal shield (815 [Ain long and 485 ^m wide) and chaetotaxy as in
female. Ventrally the idiosoma is covered by a well-ornamented holo ventral

FIGS 51-57. Hypoaspis athiasae sp. n. Male, Fig. 51. Venter. Fig. 52. Leg II. Fig.
54. Chelicera. Deutonymph, Fig. 53. Dorsal shield. Fig. 57. Venter. Proto-
nymph, Fig. 55. Dorsum. Fig. 56. Venter.

84 M. COSTA

shield, usually with 10 pairs of setae in addition to the regular anal setae (fig. 51).
The extent and outlines of the shield may vary asymetrically and with it the chaeto-
taxy. The chelicera (fig. 54) with monodentate movable digit which bears a long
slender spermadactyl, slightly curved distally. Leg II with pointed spine-like
setae on femur to tarsus (fig. 52), similar to the condition in H. krameri (Evans &
Till, op. cit.).

DEUTONYMPH : Dorsal shield (715 [xm long and 400 (xm wide) deeply incised later-
ally. Chaetotaxy similar to that of the female, central setae (mainly i series) notice-
ably long. The shield is faintly ornamented mainly on its posterior portion. The
distribution and relative lengths of the setae is shown in fig. 53. The anal shield
is rounded and the postanal seta is markedly shorter than the paranal setae (fig. 57).

PROTONYMPH: The idiosoma (530 [xm long) is covered by two dorsal shields and
three pairs of platelets in the mesonotal region. The podonotal shield (350 {xm
long and 290 (xm wide) with n pairs of long setae. The opisthonotal shield (120 pun
long and 170 (xm wide) with 8 pairs of setae. Setae 85, Z4~5 and J5 have the same
relative lengths as in the adult. The distribution and the relative lengths of the
setae are shown in fig. 55. The venter (fig. 56) with small sternal shield which has
a very indistinct anterior margin.

DIFFERENTIAL DIAGNOSIS: This species is closely related to H. rhinocerotis from
which it can be separated mainly by its longer central dorsal setae, a character
which is even more conspicuous in the deutonymph.

NOTES : This species has been collected in large numbers (by Dr C. Athias-Henriot)
from soil as well as from eggs of 0. monoceros in the Ivory Coast. It is well known
that many dynastine beetles emerge from their puparia during a few days of the
year only, usually for a short time before dusk. It may well be that the possibility
to collect a large number of the mites from the soil happened after a mass emergence
of the beetles which are probably their normal host.

MATERIAL: Holotype: $, ex humid soil (Aerodrome), Ivory Coast, July 18, 1969,
coll. C. Athias-Henriot. Paratypes: i $ and numerous female specimens ibid. ;
5 $$ Oryctes monoceros, Ivory Coast, 1969.

Hypoaspis dubius sp. n.

FEMALE: Dorsum covered by a single dorsal shield (640 [xm long and 380 {xm
wide), with 37 pairs of simple setae. The shield is finely ornamented mainly on its
posterior part. Setae 12, si and 54-6 are longer than the remaining podonotal
setae, the longest dorsal setae are Z4 (130 (xm), the shortest J5 (25 |xm). Nearly
all the central setae are long and extend beyond the base (or the horizontal level
of the bases) of the consecutive setae. The distribution and the relative lengths
of the setae are shown in fig. 58. Tectum (fig. 61) triangular with denticulate
margin.

HYPOASPIS MITES AND BEETLES

Tritosternum normal with pilose laciniae. Sternal shield (135 |xm long and 140 |j.m
wide at St2) faintly ornamented mainly in its anterior and lateral parts, posterior
margin slightly concave to nearly straight. Sti shorter than remaining sternal
setae which are long and extend beyond the base of consecutive setae. The tongue-

FIGS 58-64. Hypoaspis dubius sp. n., female. Fig. 58. Dorsal shield. Fig. 59. Venter.
Fig. 60. Leg IV. Fig. 61. Tectum. Fig. 62. Tarsus II. Fig. 63. Chelicera.
Fig. 64. Gnathosoma, ventral view.

86 M. COSTA

shaped genital shield (distance between genital setae 105 (xm) is faintly ornamented
(fig. 59). Metapodal shields narrow and elongate. The postanal seta is shorter
than the paranal setae. The peritreme extends anteriorly slightly beyond the middle
of coxa I.

Gnathosoma (fig. 64) with well-sclerotized corniculi and fimbriate inner malae,
six rows of deutosternal denticles (14-18 per row). Movable digit of chelicera
(fig. 63) bidentate, fixed digit with about 8 small denticles in addition to one larger
tooth.

The approximate lengths of the legs (excluding pretarsi) are: I 640 (xm; II
490 (Jim; III 490 {xm; IV 690 (xm. Tarsus II (fig. 62) with two dorsal distal
blunt spines. Leg IV (fig. 60) with macrosetae on the femur (145 |xm) and tarsus
only. Macrosetae are present also on femora II and III.

MALE: Unknown.

DIFFERENTIAL DIAGNOSIS: This species is characterized by its small size and
short Z4 and ad\ of femur IV.

NOTES : The status of this species is uncertain and it might be a hybrid. Originally
it was introduced from the Ivory Coast, ex Oryctes monoceros. It has become
established in W. Samoa and occurred together with H. rhinocerotis on a field-
collected 0. rhinoceros (coll. Dr M. J. Stelzer). In view of the fact that Hypoaspis
athiasae sp. n. has also been collected from 0. monoceros, a misdetermination of the
African host cannot be ruled out, since over a dozen species of Oryctes exist in central
and western Africa.

MATERIAL : Holotype : $, laboratory reared, Apia, W. Samoa, July 1969, laboratory
colony started from specimens ex Oryctes monoceros, Ivory Coast. Paratypes:
many ibid. ; 6 -$, 0. rhinoceros, Apia, W. Samoa, Sept. 1969, coll. Dr M. J. Stelzer.

Hypoaspis remilleti sp. n.

FEMALE: Dorsum covered by single dorsal shield (870 |xm long and 560 fxm
wide) leaving a wide strip of uncovered integument. The shield bears 36 pairs of
setae (53 missing, compare Hypoaspis phyllognathi sp. n.) of three different types:
minute setae (about 20 [xrn long: 13-5, Z2-3, J series px2~3, Zi-3 and Si) ; extremely
long setae (i2, 54-6 and Z4 which are 400 ^m long) and 'normal' marginal setae.
The posterior elongate integumental setae are only 180 jxm long. The distribution
and the relative lengths of the setae are shown in fig. 65. Tectum (fig. 66) triangular
with denticulate margin.

Tritosternum normal with pilose laciniae. Sternal shield (160 (xm long and 170 [xm
wide at St2) ornamented, with irregular posterior margin (fig. 67). The sternal
setae are long, extending beyond the bases of the consecutive setae. The well-
ornamented genital shield (distance between genital setae 150 (xm) expands slightly
beyond coxae IV. Anal shield with round anterior margin, postanal seta shorter
than paranal setae. The peritreme extends anteriorly to or slightly beyond the
anterior margin of coxa I, it is free both anteriorly and posteriorly.

HYPOASPIS MITES AND BEETLES

Gnathosoma (fig. 71) with well-sclerotized corniculi and fimbriate internal malae,
six rows of minute deutosternal denticles. Movable digit of chelicera (fig. 68)
bidentate, fixed digit with about 12 small denticles in addition to one larger tooth.

FIGS 65-71. Hypoaspis remilleti sp. n., female. Fig. 65. Dorsum. Fig. 66. Tectum.
Fig. 67. Venter. Fig. 68. Chelicera. Fig. 69. Tarsus II. Fig. 70. Leg IV.
Fig. 71. Gnathosoma, ventral view.

88 M. COSTA

The approximate lengths of the legs (excluding pretarsi) are: I 890 pan; II
740 pan; III 800 pan; IV noo pan. Tarsus II (fig. 69) with two blunt dorsal
subterminal spines. Leg IV (fig. 70) with macrosetae on the femur (380 pan)
and tarsus only. Leg chaetotaxy normal.

MALE : Dorsal shield as in female. The venter is covered by a holoventral shield
with 10 pairs of setae in addition to the regular anal setae. Leg II with pointed
spine-like ventral setae on femur to tarsus. Chelicera with slender spermodactyl,
distally curved.

DIFFERENTIAL DIAGNOSIS : This species can be easily recognized by its very short
central setae and its long antero-lateral setae. It differs from closely related
'Coleolaelaps' proximus Cooreman, 1948 mainly in the shape of the sternal shield
which is markedly longer than wide in C. proximus. Associated with Heteroligus
meles Billb. (Scarabaeidae : Dynastinae).

MATERIAL: Holotype: ., ex Heteroligus meles Billb., nr. Abidjan, Ivory Coast,
1969, coll. Dr M. Remillet. Paratypes: numerous specimens including males with
the same data.

Lucanaspis gen. n.

General facies of female as in Hypoaspis with the following differences: Thirty-
three pairs of dorsal setae of which i2, 54 and Z4 are very long. Sternal shield
markedly wider than long. Legs stumpy and shorter than length of dorsal shield.
Tarsus II with two subterminal stout and pointed setae.

MALE: Unknown.

Lucanaspis brachypedes sp. n.

FEMALE: Dorsum covered by a single dorsal shield (545 pan long and 360 pan
wide), devoid of ornamentation, with 33 pairs of simple setae. The podonotal
setae (17 pairs) are very short (10-15 pan) with the exception of ii and si (85 pan),
12, 54 (120 pan), 55 and s6 (60 pan). The opisthonotal setae (16 pairs) are short
with the exception of Z4 (155 pan) which are the longest dorsal setae. The distri-
bution and the relative lengths of the setae are shown in fig. 72. Tectum (fig. 78)
triangular with denticulate margins.

Tritosternum normal with pilose laciniae. Sternal shield (75 pan long and 130 pan
wide at St2) with distinct anterior concave margin and slightly irregular concave
posterior margin. The shield is devoid of distinct ornamentation. The first sternal
pores are very large and close to each other. Metasternal setae inserted on the
integument, the associated pore may be incorporated into the sternal shield (fig. 73).
Genital shield (distance between genital setae 95 pan) broad and tongue-shaped.
Anterior margin of anal shield semicircular, paranal setae slightly longer than postanal
seta. The shield is flanked by two pairs of slightly longer and stouter ventral
setae. Metapodal shields elongate, oval. The peritreme extends anteriorly
nearly to the anterior margin of coxa I, it is free both anteriorly and posteriorly.

HYPOASPIS MITES AND BEETLES

Gnathosoma (fig. 75) with well-sclerotized corniculi and fimbriate internal malae,
with six rows of tiny deutosternal denticles. Movable digit of the chelicera (fig. 74)
bidentate, fixed digit with three medium sized distal teeth and about seven small
proximal denticles.

FIGS 72-78. Lucanaspis brachypedes gen. n., sp. n., female. Fig. 72. Dorsal shield.
Fig- 73- Venter. Fig. 74. Chelicera. Fig. 75. Gnathosoma, ventral view. Fig.
76. Tarsus II. Fig. 77. Leg IV. Fig. 78. Tectum.

go M. COSTA

The approximate lengths of the legs (excluding pretarsi) are: I 520 (xm; II
450 (jim; III 440 (Jim; IV 540 (Jim. Leg chaetotaxy normal as in free-living
laelapids. Macrosetae are present on femora II-IV (on femur IV 165 [xm) and
tarsus IV (fig. 77). Tarsus II (fig. 76) with two subterminal pointed stout setae.

MATERIAL: Holotype: <j>, ex lucanid beetle, nr. Abidjan, Ivory Coast, Oct. 1969,
coll. Dr M. Remillet. Paratypes: 8 $?, same data.

Dynastaspis gen. n.

General facies of female as in Hypoaspis with the following differences: 32 pairs
of dorsal setae (18 podonotal and 14 opisthonotal) with 54-6 and Z^ very long and
'wavy'. Tectum with nondenticulate margin. Four macrosetae of a different
homology on tarsus IV: adz, alz, Ph and ^^3 (instead oipdz, pd$ and adz in Hypoaspis
s. str.), pdz being proximally inserted to adz (as distinct from being at the same level
in Hypoaspis s. str.). Tarsus II with two subterminal pointed stout setae. Orna-
mentation of genital shield different from that found in Hypoaspis.

MALE: Unknown.

Dynastaspis walhallae sp. n.

FEMALE : A single dorsal shield (1030 (xm long and 580 jxm wide) covers the dorsum
incompletely, faintly ornamented with a scale-like pattern (fig. 86), with 32 pairs
of simple setae (fig. 79). Eighteen pairs of podonotal setae, Z3 absent, 57 on the
integument and 14 pairs of opisthonotal setae (84 and px2~3 absent). Medial
setae mainly short (e.g. 14 45 (Jim), 54-6 long and 'wavy', Z^ being the longest
dorsal setae (400 (xm). Tectum (fig. 83) with nondenticulate margin and broad
median projection.

Tritosternum normal with 'pilose laciniae. Sternal shield (140 (jim long and
195 (xm wide at St2), well ornamented with concave anterior margin and deeply
concave posterior margin (fig. 80, 8ia-c). Sternal setae long, St2 extending beyond
posterior margin of shield. The shape of the sternal shield is variable, mainly
in the lateral outlines (figs 8ia-c, in 8ic one Sti completely missing). Genital
shield (distance between genital setae 155 [xm) tongue-shaped, with very distinct
ornamentation which is completely different from that found in species of Hypoaspis
s. str. Metapodal shields small and irregular, several small platelets present at
the same level. Anal shield nearly triangular in shape, postanal seta shorter than
the paranal setae. The peritreme extends anteriorly slightly beyond the posterior
margin of coxa I, it is free anteriorly and posteriorly.

Gnathosoma (fig. 87) with well-sclerotized corniculi and fimbriate internal malae,
with six rows of deutosternal teeth (10-14 P er row). Movable digit of the chelicera
(fig. 85) bidentate, fixed digit with about eight denticles of different sizes and shapes.

The approximate lengths of the legs (excluding pretarsi) are: I 1010 jxm; II
850 (xm; III 870 [xm; IV 1190 jxm. Tarsus II (fig. 84) with pointed setae only,
leg IV (fig. 82) with macrosetae on femur (400 (xm) and four macrosetae (adz, <*h,

HYPOASPIS MITES AND BEETLES 91

plz, pdz) on the tarsus with pdz proximally inserted. Otherwise leg chaetotaxy
normal.

FIGS 79-87. Dynastaspis walhallae gen. n., sp. n., female. Fig. 79. Dorsum. Fig. 80.
Venter. Fig. 8ia-c. Variations in shape of sternal shield. Fig. 82. Leg IV. Fig.
83. Tectum. Fig. 84. Tarsus II. Fig. 85. Chelicera. Fig. 86. Ornamentation of
dorsal shield. Fig. 87. Gnathosoma, ventral view.

92 M. COSTA

MATERIAL: Holotype: -, ex larvae of Dynastes tytius Linn. (Scarabaeidae :
Dynastinae), dead Black Locust Tree, Walhalla, South Carolina, August 1969,
coll. M. Palmer. Paratypes: 8 .-, same data as type.

NOTES: The occurrence of this species on larvae of Dynastes tytius is rather sur-
prising as Sikora (1968) reported a different species of 'Coleolaelaps' from the adult
of D. tytius. It may be that the host association of D. walhallae is accidental, the
true host being another woodboring beetle.

Angosomaspis gen. n.

FEMALE: Single dorsal shield with a large number (over 100) of minute (15-
20 (jun) setae, ventral integument with about 30 pairs of long, very attenuated
whip-like setae. Macroseta of femur IV exceedingly long, extending beyond middle
of tarsus IV, only one macroseta (pd%) on tarsus IV. Long macroseta also on femora
II and III. Tarsus II with two subterminal spine-like setae which are originally
pointed, though usually the tip is broken off, leaving a sharp (not blunt) end.
Fixed digit of chelicera with about a dozen sharp, needle-shaped denticles proximal
to the pilus dentilis. Remaining characters similar to Hypoaspis.

MALE: Unknown.

Angosomaspis multisetosus sp. n.

FEMALE: Dorsum covered by single dorsal shield (800 (jon long and 480 (xm
wide) covering most of the dorsum. Over a 100 minute setae (15-20 (jun) are distri-
buted on the shield, obscuring any paired arrangement. Only four pairs of anterior
setae and one pair of posterior marginal setae are of 'normal' Hypoaspis-type
length and appearance (60-95 [Am). The distribution and the relative lengths of
the setae are shown in fig. 88. The laterodorsal chaetotaxy is obscured by a row
of setae which are inserted on the soft integument between the narrow peritrematal
shield and the dorsal shield, in mounted specimens this part of the integument in-
variably folds below the dorsal shield and only the examination of the mite in lateral
position reveals that these setae are not inserted on the shield. The dorsal shield
is finely granulated but shows no ornamentation. Tectum (fig. 91) triangulate
with margins partly or totally denticulate.

Tritosternum normal with long pilose laciniae. Sternal shield (190 {xm long
and 170 fxm wide at St2) with straight anterior and posterior margin, finely granu-
lated surface without ornamentation. Sternal setae long (e.g. St2 150 ^m) and
whiplike, very attenuated and coming to a fine end, they are similar to the remaining
(about 30 pairs) ventral setae. Genital shield large and elongate (distance between
genital setae 145 (Am), devoid of ornamentation. Anal shield with slightly curved
anterior margin, paranal setae longer than postanal seta. The peritreme extends
anteriorly slightly beyond the middle of coxa I, it is accompanied by a very narrow
external and internal peritrematal shield which is free both anteriorly and posteriorly.

HYPOASPIS MITES AND BEETLES

Gnathosoma (fig. 94) with well-sclerotized corniculi and fimbriate internal
malae with six rows of minute deutosternal denticles. Movable digit of chelicera
(fig. 92) bidentate, fixed digit with about a dozen fine pointed denticles proximal to
the pilus dentilis.

FIGS 88-94. Angosomaspis multisetosus gen. n., sp. n., female. Fig. 88. Dorsum. Fig. 89.
Venter. Fig. 90. Leg IV. Fig. 91. Tectum. Fig. 92. Chelicera. Fig. 93. Tarsus
II. Fig. 94. Gnathosoma, ventral view.

94 M. COSTA

The approximate lengths of the legs (excluding pretarsi) are: I 760 (xm; II
650 (j.m; III 680 (Jim; IV 890 [xm. Long macrosetae on femora II-IV, ad\ of
femur IV (490 [xm) extending beyond the middle of the corresponding tarsus.
Only one macroseta (pdz) on tarsus IV (fig. 90). Tarsus II (fig. 93) with two sub-
terminal pointed spine-like setae, usually the tip is broken off leaving a sharp
jagged surface.

MATERIAL: Holotype: $, ex Angosoma centaurus (Scarabaeidae : Dynastinae),
nr. Abidjan, Ivory Coast, 1969, coll. Dr M. Remillet. Paratypes: 20 $?, same
data as type.

Promacrolaelaps gen. n.

FEMALE : Large mites with a convex dorsal shield which covers the sides and the
dorsum. Thirty-one pairs (17 podonotal and 14 opisthonotal) of long setae are
inserted on the shield. Genital shield large, expanded posterior to the genital setae.
Macrosetae present on femora I-IV, genua III-IV and single macroseta on tarsus IV.
Tarsus II without spine-like or spur-like setae. Gnathosoma with seven rows of
deutosternal denticles, fixed digit of chelicera with six medium-sized sharp teeth
proximal to the pilus dentilis.

MALE: Unknown.

Promacrolaelaps hunteri sp. n.

FEMALE: The dorsum is completely covered by a large convex dorsal shield
(1260 |xm long and 820 [xm wide) which covers the mite also laterally. Thirty-one
pairs of simple long setae (17 podonotal and 14 opisthonotal) are inserted on the
shield. Setae Z4 are the longest (510 [xm) dorsal setae. The shape of the shield
as well as the lateral displacement of many setae did not permit positional homo-
logization of most setae, their distribution and relative lengths being shown in
fig. 95. Tectum (fig. 97) with denticulate margin.

Tritosternum normal, with pilose laciniae. Sternal shield (170 (xm long and 210 [xm
wide at St2) wider than long, with long sternal setae extending considerably beyond
the bases of consecutive setae. The shield is ornamented and granulate, its anterior
and posterior margins are straight to slightly concave. Genital shield (fig. 96)
large (distance between genital setae 220 (xm), well ornamented and expanding
posterior to the genital setae. Metapodal shields small and irregular. Anal shield
with rounded anterior margin, postanal seta slightly longer than paranal setae.
The anal shield is flanked by a pair of long (370 [xm) setae. Approximately 17 pairs
of setae are inserted on the integument posterior to coxae IV. The peritreme
extends anteriorly beyond the middle of coxa I and is free both anteriorly and
posteriorly.

Gnathosoma (fig. 100) with well-sclerotized corniculi and fimbriate internal
malae, seven rows of deutosternal teeth (10-20 per row) are present. Movable
digit of the chelicera (fig. 99) bidentate, fixed digit with six well-defined, medium-
sized sharp teeth proximal to the pilus dentilis.

HYPOASPIS MITES AND BEETLES

The approximate lengths of the legs (excluding pretarsi) are: I 980 (xm; II
850 fxm; III 830 (Jim; IV 930 [xm. Macrosetae present in femora I and II
(pdi), III and IV (ad\, 450 (j.m long on femur IV) ; on genua III and IV (ad\, 450 (xm
long on genu IV) and a single macroseta (pdz 250 yjri) on tarsus IV (fig. 101).

'lil/'MN '' ' '' 1 ' N

; KiVV.M v ' III >

100

FIGS 95-101. Promacrolaelaps hunteri gen. n., sp. n., female. Fig. 95. Dorsal shield.
Fig. 96. Venter. Fig. 97. Tectum. Fig. 98. Tarsus II. Fig. 99. Chelicera.
Fig. 100. Gnathosoma, ventral view. Fig. 101. Leg IV.

96 M. COSTA

Tarsus II (fig. 98) with simple pointed setae only, the ventral setae being slightly
stouter.

This species is named in honour of my friend and colleague Dr P. E. Hunter
(Department of Entomology, The University of Georgia).

MATERIAL: Holotype: 9 ex Promacrus bimucronatus Pallas (Scarabaeidae :
Euchirinae), Tivon, Israel, August 20, 1967, coll. M. Costa. Paratypes: 5 $<j>,
same data as type.

DISCUSSION

The vague early definition of genera and species can often be applied to several
different taxa subsequently discovered. In the present study the early descriptions
and figures of Hypoaspis krameri Can. apply equally well to H. krameri Can. sensu
Evans & Till (1966), H. neokrameri sp. n. ; H. pentodoni sp. n. ; H . phyllognathi sp. n. ;
H. integer Berlese sensu Samsinak (1960), etc. Vitzthum (1940-43) records 'H.
krameri' from the following hosts: 0. nasicornis, Cetonia aurata, Potosia floricola,
Pentodon punctatus, Polyphylla fullo apparently placing under H. krameri several
species with a higher degree of host specificity than assumed by him. The un-
certainty of the determination of mites of the Hypoaspis and Coleolaelaps com-
plexes by Berlese has been discussed by Costa & Hunter (op. cit.). In view of this
early species confusion and the fact that their material was obtained from a new
host, namely Lucanus sp., it seems rather uncertain that H. krameri Can. sensu
Evans & Till is actually conspecific with the type species, it undoubtedly agrees
with the description and definition of Hypoaspis s. str. However, in order to avoid
additional confusion on the subject, the decision made by Evans & Till is accepted
here until the type (? lost) or topotypic material from the type host can be examined.
The definition of Hypoaspis s. str. as conceived here, as well as the definition of
Coleolaelaps Berl. as conceived by Costa & Hunter (op. cit.) will necessitate the
transfer of many species from the latter genus to Hypoaspis s. str., this being,
however, outside the scope of the present study.

Being convinced that host-association is just as good a character for taxonomic
discrimination as morphological characters, I have designated four new genera
of which the first two (Lucanaspis and Dynastaspis) are closer related to Hypoaspis
s. str. than the remaining two (Angosomaspis and Promacrolaelaps) . As additional
species are expected for these genera, the generic definitions are short and not very
rigorous, allowing amendment for the inclusion of new species. The generic
delimitations being to a large degree a matter of personal opinion, I prefer this
treatment to the lumping of many different forms for 'phylogenetic' reasons.

The juvenile forms of Hypoaspis s. str. are described here for the first time (H.
rhinocerotis and H. athasiae) and they agree to the general type of the free-living
laelapids (vide H. aculeifer (Can.), Evans & Till, op. cit.). I should like to point
out the 'long-haired' nature of the juveniles: the protonymph having longer setae
than the deutonymph and this having longer setae than the adult.

The occurrence of macrosetae on the idiosoma as well as on the legs, mainly
leg IV, seems to have arisen independently in various mesostigmatic mites associated

HYPOASPIS MITES AND BEETLES 97

with arthropods. The function of the macrosetae is at present unknown, it may
be mainly thigmotactic and concerned with finding the right location on the host,
or it may also serve to avoid overcrowding on the host, assuring each mite its own
volume of space.

SUMMARY

The following species and genera of mites associated with phytophagous lamellicorn
beetles are described and figured: Hypoaspis neokrameri sp. n. ($); Hypoaspis
pentodoni sp. n. ($, <) ; Hypoaspis phyllognathi sp. n. ($) ; Hypoaspis integer Berlese
($) ; Hypoaspis rhinocerotis Ouds. ($, $, dn, pn) ; Hypoaspis athiasae sp. n. ($, <,
dn, pn) ; Hypoaspis dubius sp. n. ($) ; Hypoaspis remilleti sp. n. (?) ; Lucanaspis
brachypedes gen. n., sp. n. ($) ; Dynastaspis walhallae gen. n., sp. n. ($) ; Angosomaspis
multisetosus gen. n., sp. n. (?) ; Promacrolaelaps hunteri gen. n., sp. n. ($).

ACKNOWLEDGEMENTS

I would like to thank the following for the loan or presentation of material used
in this study: Dr C. Athias-Henriot (Laboratoire du Sol de I.N.R.A., Dijon); Dr
Lee H. Herman, Jr. (American Museum of Natural History, New York); Dr M.
Remillet (O.R.S.T.O.M., Centre d'Adiopodoume, Abidjan, Ivory Coast); Dr K.
Samsinak (Entomological Institute of the Czechoslovak Akademy of Sciences,
Prague) ; Dr M. J. Stelzer and Mr B. Zelazny (U.N./S.P.C. Rhinoceros Beetle Project,
Apia, Western Samoa). I am grateful to Dr F. Pegazzano (Stazione di Entomo-
logia Agraria, Florence) for information concerning specimens in the Berlese col-
lection.

It is my pleasure to express greatful thanks to Dr P. E. Hunter (Department
of Entomology, The University of Georgia, Athens) for his generous hospitality,
his continuous interest and for his criticism of parts of the manuscript.

This study was made while holding a Senior Foreign Scientist Fellowship of the
National Science Foundation at the Department of Entomology, The University
of Georgia.

REFERENCES

COSTA, M. & HUNTER, P. E. (in press). The genus Coleolaelaps Berlese, 1914 (Acarina: Meso-
stigmata) .

EVANS, G. O. 1958. A Review of the Laelaptid Paraphages of the Myriapoda with descriptions

of three new species (Acarina: Laelaptidae) . Parasitology 45 : 352-368.

1963. Observations on the chaetotaxy of the legs in the free-living Gamasina (Acasi :
Mesostigmata) . Bull. Brit. Mus. nat. Hist. (Zool.). 10 (5) : 275-303.

EVANS, G. O. & TILL, W. M. 1966. Studies on the British Dermanyssidae (Acari: Mesostig-
mata). Pt II Classification. Bull. Brit. Mus. nat. Hist. (Zool.). 14 (5) : 109-370.

GRANDI, G. 1925. Contributo alia conoscenze biologica e morphologica di alcuni Lamellicorni
fillifagi e descrizione di una nuova specie di Acaro. Boll. Lab. Zool. gen. agr. R. Scuola
Agric. Portici 18 : 159-224.

98 M. COSTA

HUNTER, P. E. & COSTA, M. (in press). Gymnolaelaps shealsi n. sp. (Acarina: Mesostigmata)

associated with the imported fire ant.
SAMSINAK, K. 1960. Kurze Bemerkungen uber Mesostigmata (Acari). Cas. 6sl. Spol. ent.

57 (3) : 275-284.
SIKORA, W. B. 1968. A review of the genus Coleolaelaps Berlese, 1914 with descriptions of

three new species (Acarina: Mesostigmata). M.S. Thesis, University of Georgia, Athens,

Georgia, U.S.A. pp. 43.
VITZTHUM, H. 1940-43. Acarina in H. G. Bronn (ed.), Klassen und Ordnungen des Tierreiches.

Vol. 5, Part 5, Book 5, ion pp., Leipzig.

Dr M. COSTA, M.Sc., Ph.D.
KIBBUTZ MISHMAR HAEMEK
ISRAEL

A LIST OF SUPPLEMENTS
TO THE ZOOLOGICAL SERIES

OF THE BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

1. KAY, E. ALISON. Marine Molluscs in the Cuming Collection British Museum
(Natural History) described by William Harper Pease. Pp. 96; 14 Plates.
1965. (Out of Print.) 3.75.

2. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier and
Valenciennes. Pp. 180; n Plates, 15 Text-figures. 1967. 4.

3. TAYLOR, J. D., KENNEDY, W. J. & HALL, A. The Shell Structure of Mineralogy
at the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125; 29 Plates,
77 Text-figures. 1969. 4.50.

4. HAYNES, J. R. Cardigan Bay recent Foraminifera (Cruises of the R.V. Antur)
1962-1964. (In press.)

Printed in England by Staples Printers Limited at their Kettering, Northants, establishment

PI97J

THE POLYCHAETE FAUNA OF THE
SOLOMON ISLANDS

P. E. GIBBS

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 21 No. 5

LONDON : 1971

THE POLYCHAETE FAUNA OF THE
SOLOMON ISLANDS

PETER EDWIN GIBBS

Marine Biological Association of the U.K., Plymouth

Pp. 99-211; 17 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)

ZOOLOGY Vol. 21 No. 5

LONDON: 1971

THE BULLETIN OF THE BRITISH MUSEUM

(NATURAL HISTORY), instituted in 1949, is issued
in five series corresponding to the Departments of the
Museum, and an Historical series.

Parts will appear at irregular intervals as they
become ready. Volumes will contain about three or
four hundred pages, and will not necessarily be completed
within one calendar year.

In 1965 a separate supplementary series of longer
papers was instituted, numbered serially for each
Department.

This paper is Vol. 21, No. 5 of the Zoological series.
The abbreviated titles of periodicals cited follow those
of the World List of Scientific Periodicals.

World List abbreviation
Bull. Br. Mus. not. Hist. (Zool.).

Trustees of the British Museum (Natural History), 1971

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued 21 September, 1971 Price 3-55

THE POLYCHAETE FAUNA OF THE
SOLOMON ISLANDS

By P. E. GIBBS

CONTENTS

Page

SYNOPSIS ........... ioi

INTRODUCTION ........ . ioi

GENERAL ACCOUNT .......... 102

COMPOSITION OF THE POLYCHAETE FAUNA AND ITS ZOOGEOGRAPHICAL

AFFINITIES .......... 104

ECOLOGICAL OBSERVATIONS

Littoral survey . . . . . . . . . in

Dredge survey of Marovo Lagoon . . . . . . 114

Commensal polychaetes . . . . . . . . 116

Brackish water and terrestrial species . . . . . 117

Spawning of Eunice (Palola) siciliensis . . . . . 118

SYSTEMATIC ACCOUNT . . . . . . . . . 119

REFERENCES ........... 205

SYNOPSIS

An account of the polychaete fauna of the Solomon Islands (western Pacific Ocean) is given,
based on a large collection which was obtained during the Royal Society Expedition in 1965. A
total of 220 species are recorded. Two new genera (Family Nereidae), 13 new species and 3 new
subspecies are described. The zoogeographical affinities of the fauna are analysed and certain
ecological aspects discussed.

INTRODUCTION

As pointed out by Knox (1958), the polychaetes of the Pacific Islands are poorly
known and much work remains to be done before a relatively complete bio-
geographical analysis can be made. Apart from reports on small collections or
individual families, comprehensive accounts of the polychaetes from any of the
island groups in the tropical west Pacific regions are few. Augener (19270) records
47 species from New Britain and Fauvel (1947) describes about 100 species from
New Caledonia. The fauna of the Marshall Islands is comparatively well known
with 131 species recorded as a result of the work of Hartman (1954) and, more
recently, of Reish (1968).

The Solomon Islands are situated between the latitudes of 5S and 12 S and
between the longitudes of 155 E and 163 E. From the zoogeographical viewpoint
these islands are of great interest since they lie between the faunistic centre of the

102 P. E. GIBBS

Indo-west-Pacific region, the Malay Archipelago, and the more distant islands of
the Central Pacific. Thus, one of the primary aims of the Expedition organized
by the Royal Society of London in 1965 was to determine the biogeographical
relationships of these islands with the adjacent regions. As a member of the Marine
Party of this Expedition the author spent five months, from July to December,
investigating the polychaete fauna and this paper is an account of the ecological
and faunistic observations that were made during this period.

I am indebted to the Royal Society, Professor E. J. H. Corner, F.R.S., (Expedition
Leader) and Professor J. E. Morton (Leader of the Marine Party) for the opportunity
to participate in the Expedition to the Solomon Islands in 1965. For their help in
the field I have to thank the members of the Marine Party, particularly Dr. D. R.
Stoddart, Captain S. B. Brown and crew of the A. K. Maroro, and our native
assistants, especially Ini Munamaori. I am also grateful to Wilson Ifunaoa for
providing a valuable collection of the 'palolo' rising from Fanalei, Malaita.

For their kind assistance and helpful advice in the identification of various
specimens and families, I have to thank Dr. Olga Hartman (two new genera of
Nereidae), Dr. Minoru Imajima (Syllidae), Dr. Charlotte Mangum (Maldanidae) and
Professor G. P. Wells (Abarenicola) . Other groups were identified by Dr. P.
Bergquist (Porifera), Miss A. Clark (Echinodermata) and Dr. S. J. Edmonds
(Sipunculoidea). I am also indebted to Dr. J. D. George for the loan of specimens
from the collections of the British Museum (Natural History) .

GENERAL ACCOUNT

The Marine Party of the Expedition was based and carried out investigations on
Guadalcanal, in the Florida Islands (Nggela Group), Russell Islands and New
Georgia Group (for a detailed account of the Expedition, see Corner, 1969). In these
areas, collections of polychaetes were obtained from the following localities (see
figs i and 2) : western end of Guadalcanal - Komimbo Bay, Cape Esperance and
Naro Bay ; north coast of Guadalcanal - Mamara Point, Honiara and Lunga Point ;
eastern end of Guadalcanal (Marau Sound) - Graham Point, Fintry Point, Marauni-
bina Island, Pigeon Island, and Lauvie Island ; Florida Islands - Kokomtambu
Island, Tetel (or Gaskell) Island, Nggela (or Florida) Island and Haroro; Russell
Islands (Banika Island) - Yandina, Sifola and Lingatu ; New Georgia Group (Marovo
Lagoon) - Matiu Island, Pirikale Island, Paleki Island and Batuona (or Wickham)
Island. Additional collections were made at Gizo Island (New Manra) and on
Malaita - Auki Harbour, Kalota Island, Alite Harbour and at the southern end of
Maramasike Passage.

The polychaete fauna of the moderate to very exposed coral reef platforms
consists chiefly of coral-boring and crevice-dwelling species. To obtain such species
it is necessary to break open the coral rock with hammer and chisel and then care-
fully extract the worms from their burrows or from crevices using forceps. Those
species living amongst the algal cover or chaetopterid tubes are best obtained by
dissecting and then washing these materials. Investigations of these habitats were

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS

103

FIG. i. Map of the Solomon Islands showing collection localities mentioned in the text.
Insets - Marau Sound (left) and parts of the Russell Islands and Florida Islands (right).

io 4 P. E. GIBBS

made at (collection dates in brackets) Mamara Pt. (24.viii), Sifola (2.ix), Lingatu
(19. x), Matiu and Paleki Is. (3-7.vii and 27-3i.viii), Batuona I. (3-ix) and at New
Manra (4-5 .ix). Further collections were made at Kokomtambu I. (2.vii) and at
Maramasike Pg. (26.xi). Beachrock fauna was investigated at Lauvie I. (iS.ix and
4 .x).

On the more sheltered shorelines a wide range of sedimentary deposits was studied.
The larger species of the infauna were obtained by digging and the smaller species
by sieving samples of the deposit through 0-5, i-o or 2-0 mm mesh, the mesh size
depending on the grade of the sediment. Investigations of the infauna inhabiting
a variety of deposits, ranging from thick mud to shell gravel and coral debris, were
carried out at Tetel I., Haroro and Nggela I. (5-io.vii and 24-2g.vii), Komimbo Bay,
Cape Esperance and Naro Bay (i8-2O.vii and i6-2i.viii), Graham Pt., Fintry Pt.
and Maraunibina I. (i7.ix-io.x). Small collections were also made at Yandina
(i8.x) and Honiara (28. xi).

In addition to the above studies, further species were obtained from a brackish
water lagoon at Lunga Point near the mouth of the Lunga River (g.ix-n.ix) and
from encrusting organisms, particularly sponges, on wharf piles at Yandina (i9-20.x
and 5.xi). At a number of the localities mentioned above and including Auki Hr.
and Kalota I. (20-22. xi) commensal polychaetes were discovered. It was not
possible for the author to be present at a rising of the 'palolo' but a small collection
of the spawning that took place at Fanalei on 2~3.xi.66 was kindly forwarded by
Wilson Ifunaoa. Dredging operations were confined to the New Georgia Group
where a survey of the benthos of Marovo Lagoon was carried out during the period
22.x-i4.ix in conjunction with the studies of Dr. D. R. Stoddart.

The nomenclature used in this paper is that of Hartman (1959, 1965) although
later revisions by various authors have been followed. The classification used by
Day (1967) has been adopted. The usage of the place-names in the Solomon
Islands is that given on the Directorate of Overseas Survey Map (i : 1,000,000).
Other names are taken from 'the Admiralty Charts or are local names, the spelling
of which must be regarded as provisional.

The collections, including the type specimens, have been deposited in the British
Museum (Natural History).

COMPOSITION OF THE POLYCHAETE FAUNA AND ITS
ZOOGEOGRAPHICAL AFFINITIES

The collection of polychaetes from the Solomon Islands comprises over 2000
specimens and is composed of 220 species.* Of this total, 88 (40%) of the species are
represented by only one or two specimens. Thus the actual number of species in
the fauna must be much higher, perhaps as high as 400, since probably many of the
smaller and less conspicuous species, including many of the syllids as well as the
paraonids and spirorbids, were overlooked in the field. The species composition of
the fauna by families is given in Table i. The errantiate families Aphroditidae
(24 species), Eunicidae (22) and Syllidae (20) are well represented and in total account

* See Appendix

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS

105

for about one-half of the errant species. Sedentary species are fewer (97 out of the
total of 220) with the family Spionidae (15) containing the most species. In the

TABLE i

The species composition of the Solomon Islands fauna by families.

Family

No. of
species

Aphroditidae 24

Palmyridae 3

Amphinomidae 10

Phyllodocidae 13

Pilargidae 2

Hesionidae 4

Syllidae 20

Sphaerodoridae i

Nereidae 16

Nephtyidae 3

Lacydoniidae i

Glyceridae 4

Eunicidae 22

Orbiniidae 2

Spionidae 15

Magelonidae 2

Trochochaetidae 2

TOTAL

Family No. of

species

Chaetopteridae
Cirratulidae
Cossuridae

5
i

Flabelligeridae
Scalibregmidae
Opheliidae
Sternaspidae
Capitellidae
Arenicolidae

Maldanidae

Oweniidae
Sabellariidae

Pectinariidae

Ampharetidae
Terebellidae

2
IO

Sabellidae

Serpulidae

220

collection, two new genera, namely Cryptonereis n.gen. and Solomononereis n.gen.
(Family Nereidae), 13 new species and three new subspecies are present. The new
species and subspecies are the following:

Family Aphroditidae

Family Hesionidae
Family Nereidae

Family Nephtyidae
Family Eunicidae
Family Spionidae

Family Trochochaetidae
Family Flabelligeridae
Family Opheliidae
Family Oweniidae
Family Terebellidae

Hololepidella ophiuricola n.sp.
Lepidasthenia guadalcanalis n.sp.
Gyptis maraunibinae n.sp.
Cryptonereis malaitae n.gen., n.sp.
Solomononereis marauensis n.gen., n.sp.
Nephtys (Aglaophamus) munamaorii n.sp.
Eunice marovoi n.sp.
Dispio maroroi n.sp.
Polydorella novaegeorgiae n.sp.
Prionospio tetelensis n.sp.
Scolelepis squamata mendanai n. subsp.
Poecilochaetus serpens honiarae n. subsp.
Diplocirrus glaucus orientalis n. subsp.
Ophelia koloana n.sp.
Myriochele heruensis n.sp.
Pista dibranchis n.sp.

io6

P. E. GIBBS

The species collected in the Solomon Islands are listed in Table 2. From the
material three species - all maldanids - can be identified to the subfamily level only,
17 are referable to their genus and a further six can be given near identifications to
comparable species. Excluding these species and also the new species and sub-
species a total of 178 species is available for analysis.

TABLE 2

List of species taken in the Solomon Islands.

APHRODITIDAE

Gastrolepidia clavigera Schmarda

Harmothoe nigricans Horst

Harmothoe sp.

Hololepidella nigropunctata (Horst)

H . ophiuricola n.sp.

Iphione muricata (Savigny)

Lepidasthenia elegans (Grube)

L. maculata Potts

L. guadalcanalis n.sp.

L. microlepis Potts

L. stylolepis Willey

Lepidonotus (Thormora) jukesi (Baird)

Paradyte crinoidicola (Potts)

Paralepidonotus ampulliferus (Grube)

P. indicus (Potts)

Polyodontes maxillosus (Ranzani)

P. melanonotus (Grube)

Psammolyce zeylanica Willey

Sigalion bandaensis Horst

Sthenelais heterochela Horst

S. zeylanica Willey

Sthenelanella ehlersi (Horst)

Sthenolepis japonica (Mclntosh)

Thalenessa digitata Mclntosh

PALMYRIDAE

Bhawania goodei Webster
B. pottsiana Horst
Paleanotus debilis (Grube)

AMPHINOMIDAE

Amphinome nigrobranchiata Horst

Chloeia conspicua Horst

Euphrosine foliosa Aud. & M.-Ed.

E. myrtosa Savigny

Eurythoe complanata (Pallas)

? Eurythoe sp.

Notopygos gregoryi Holly

N. sibogae Horst

Pareurythoe pitipanaensis De Silva

Pseudeurytlwe paucibranchiata Fauvel

PHYLLODOCIDAE

Eteone japanensis Mclntosh

Eulalia albopicta Marenzeller

E. viridis (Linnaeus)

E. (Pterocirrus) magalhaensis Kinberg

Notophyllum splendens (Schmarda)

Phyllodoce fristedti Bergstrom

P. malmgreni Gravier

P. pruvoti Fauvel

P. quadraticeps Grube

P. (Anaitides) madeirensis Langerhans

P. (Anaitides) parva Hartmann-SchrSder

P. (Genetyllis) castanea (Marenzeller)

P. (Genetyllis) gracilis Kinberg

PILARGIDAE

Sigambra hanaokai (Kitamori)
Synelmis albini (Langerhans)

HESIONIDAE

Gyptis capensis (Day)
G. maraunibinae n.sp.
Hesione splendida Savigny
Leocrates chinensis Kinberg

SYLLIDAE

Autolytus sp.

Brania clavata (Claparede)
Exogone gemmifera Pagenstecher
E. uniformis Hartman
E. verugera (Claparede)
Haplosyllis spongicola (Grube)
Langerhansia cornuta (Rathke)
L.? rosea (Langerhans)
Sphaerosyllis hirsuta Ehlers
Syllis longissima Gravier
Syllis sp. cf. gracilis Grube
Trypanosyllis coeliaca Claparede
T. (Trypanedenta) sp.
Typosyllis alternata (Moore)
T. armillaris (Miiller)
T. brachycola (Ehlers)

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS

107

T. exilis (Gravier)
T. lucida (Chamberlin)
T. prolifera (Krohn)
Ty posy His sp.

SPHAERODORIDAE

Sphaerodoridium claparedii (Greeff)

NEREIDAE

Ceratonereis costae (Grube)

C. erythraeensis Fauvel

C. mirabilis Kinberg

Cryptonereis malaitae n.g.n.sp.

Namalycastis indica (Southern)

Nereis (Neanthes} caudata (Delle Chiaje)

N. (Neanthes) unifasciata Willey

N. (Neanthes} cf. kerguelensis Mclntosh

Perinereis cultrifera (Grube)

P. nigropunctata (Horst)

P. nuntia (Savigny)

Platynereis insolita Gravier

Pseudonereis anomala Gravier

P. masalacensis (Grube)

P. variegata (Grube)

Solomononereis marauensis n.g.n.sp.

NEPHTYIDAE

Nephtys (Aglaophamus) munamaorii n.sp.
N. (Micronephthys} sphaerocirrataWesenberg-

Lund
N. (Nephtys) sp. cf. palatii Gravier

LACYDONIIDAE

Paralacydonia weberi Horst

GLYCERIDAE

Glycera gigantea Quatrefages
G. lancadivae Schmarda
G. longipinnis Grube
G. rouxi Aud. & M.-Ed.

EUNICIDAE

Arabella tricolor (Montagu)

A. mutans (Chamberlin)

Dorvillea sp.

Drilonereis major Crossland

Eunice afra Peters

E. antennata (Savigny)

E. aphroditois (Pallas)

E. coccinea Grube

E. grubei Gravier

E. marovoi n.sp.

E. norvegica (Linnaeus)

E. tentaculata Quatrefages

E. tubifex Crossland

E. (Palola) siciliensis Grube

Lumbrineris latreilli Aud. & M.-Ed.

L. papillifera (Fauvel)

L. sphaerocephala (Schmarda)

Lysidice collaris Grube

Marphysa macinloshi Crossland

Nematonereis unicornis (Grube)

Oenone fulgida (Savigny)

Onuphis (Nothria) holobranchiata Marenzeller

ORBINIIDAE

Haploscoloplos bifurcatus Hartman
Naineris laevigata (Grube)

SPIONIDAE

Dispio maroroi n.sp.

Laonice cirrata (Sars)

Malacoceros indicus (Fauvel)

Nerinides sp. cf. gilchristi Day

Polydorella novaegeorgiae n.sp.

Prionospio cirri/era Wir6n

P. ehlersi Fauvel

P. malmgreni Claparede

P. pinnata Ehlers

P. steenstrupi malayensis Caullery

P. tetelensis n.sp.

Pseudopolydora corallicola Woodwick

Pseudopolydora sp.

Scolelepis squamata mendanai n.subsp.

Spio filicornis (Miiller)

MAGELONIDAE

Magelona japonica Okuda
Magelona sp.

TROCHOCHAE TIDAE

Poecilochaetus serpens honiarae n.subsp.
P. tropicus Okuda

CHAE TOP TERIDAE

Chaetopterus variopedatus (Renier)
Mesochaetopterus Sagittarius (Claparede)
Phyllochaetopterus elioti Crossland
P. herdmani (Hornell)
P. socialis Claparede
Spiochaetopterus costarum costarum
(Claparede)

CIRRATULIDAE

Cirriformia filigera (Delle Chiaje)
C. punctata (Grube)

io8

P. E. GIBBS

Dodecaceria fistulicola Ehlers
D. laddi Hartman
Tharyx sp.

COSSURIDAE

Cossura coasta Kitamori

FLABELLIGERIDAE

Diplocirrus glaucus orientalis n.subsp.

SCALIBREGMIDAE

Hyboscolex longiseta Schmarda
Scalibregma inflatum Rathke

OPHELIIDAE

Armandia lanceolata Willey
A. leptocirrus (Grube)
A. longicaudata (Caullery)
Ophelia koloana n.sp.
Polyophthalmus pictus (Dujardin)

STERN ASPIDAE

Sternaspis scutata (Renier)

CAPITELLIDAE

Capitellethus dispar (Ehlers)
Capitobranchus sp.
Dasybranchus caducus (Grube)
Mastobranchus dollfusi Fauvel
M. trinchesii Eisig
Mediomastus sp. cf. capensis Day
Notomastus sp.

ARENICOLIDAE

Abarenicola claparedii claparedii (Levinsen)

MALDANIDAE

Clymenella (= Macroclymene) sp. i
Clymenella (= Macroclymene) sp. 2
Clymenella (= Euclymene) sp.
Euclymeninae sp. A.
Euclymeninae sp. B.
Euclymeninae sp. C.
? Nicomache sp.
Praxillella sp.

OWENIIDAE

Myriochele eurystoma Caullery
M. heruensis n.sp.

Myriochele sp.

Owenia fusiformis Delle Chiaje

SABELLARIIDAE

Lygdamis ehlersi (Caullery)
L. indicus Kinberg

PEC TINARHDAE

Pectinaria (Pectinaria) antipoda Schmarda

AMPHA RE TIDAE

Isolda pulchella Miiller
? Sosane wireni Caullery

TEREBELLIDAE

Amaeana trilobata (Sars)
Euthelepus kinsemboensis Augener
Eupolymnia nebulosa (Montagu)
Loimia medusa (Savigny)
Lysilla ubianensis Caullery
Pista dibranchis n.sp.
P. typha (Grube)

Reteterebella queenslandia Hartman
Terebella ehrenbergi Grube
Terebellides stroemi Sars

SABELLIDAE

Branchiomma cingulata (Grube)

Hypsicomus phaeotaenia (Schmarda)

Megalomma intermedium (Beddard)

M. linaresi (Rioja)

M. quadrioculatum (Willey)

M. trioculatum Reish

M. vesiculosum (Montagu)

Potamilla ehlersi Gravier

Sabella fusca Grube

5. melanostigma Schmarda

Sabellastarte sanctijosephi (Gravier)

SERPULIDAE

Filograna implexa Berkeley
Hydroides minax (Grube)
H. uncinata (Philippi)
Mercierella enigmatica Fauvel
Serpula hartmanae Reish
S. vermicularis Linnaeus
Spirobranchus coutierei (Gravier)
5. giganteus (Pallas)
Vermiliopsis glandigerus Gravier

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS

109

At this stage a zoogeographical analysis of the Solomon Islands fauna can be
regarded only as provisional since many areas within the Indo-Pacific still remain
to be investigated and further records, as well as taxonomic revisions, will modify
the known distribution patterns of many species. The data for this analysis are
given in Table 3. The faunistic regions outlined in this table are necessarily broad:
in the Pacific region, only those island groups for which the polychaete faunas are
fairly well known are considered. As far as possible, synonymies have been taken
into account. In compiling the distributional data for each species the following
references have been consulted :

(i) Eastern Atlantic

(ii) Mediterranean Sea
(iii) South Africa
(iv) East Africa (Red Sea to

32S)

(v) Indian waters (Persian
Gulf to Andaman Is.)

(vi) South-west Australia

(vii) Malay Archipelago (Malaya
to New Guinea and
including Philippine and
Palau Is.)

(viii) East Australia (south
to Port Jackson)

(ix) New Caledonia
(x) Central Pacific (Society
and adjacent Islands)

(xi) New Zealand

(xii) Marshall Islands
(xiii) Hawaii
(xiv) Japan

Augener, 1918; Fauvel, 1923, 1927; Kirke-

gaard, 1959.
Fauvel, 1923, 1927.
Day, 1967.
Crossland, 1903, 1904, 1924; Day, 1962, 1967;

Gravier 1900-1908.
De Silva, 1961, 1965, 19650; Fauvel, 1953;

Tampi & Rangarajan, 1964; Wesenberg-

Lund, 1949.
Augener, 1913, 1914; Fauvel, 1922; Kott,

1949.
Caullery, 1944; Ehlers, 1920; Fauvel, 1935,

1939; Horst, 1910-1924; Mesnil & Fauvel,

1939; Okuda 19370; Pillai, 1965.

Augener, 1927; Monro, 1931; Rullier, 1965;

Russell, 1962; Straughan, 1967.
Fauvel, 1947.
Fauvel, 1947; Monro, 1928, 1939, 19390.

Augener, 1924, 1926; Ehlers, 1904; Knox,

1951, 19510, 1960.
Hartman, 1954; Reish, 1968.
Hartman, 1966; Straughan, 1969.
Imajima & Hartman, 1964; Imajima, 1966-

1967.

Of the total of 178 species recorded from the Solomon Islands, 21 or n-8% are
considered to be cosmopolitan in their distribution and can be eliminated from the
analysis in order to obtain a clearer pattern of the faunal relationships.

As expected, the Solomon Islands fauna has a large overlap with the faunas of the
Malay Archipelago (61-1%) and of Indian waters (64-3%), and over one-half (55-4%)
of the species are known to extend to the east coast of Africa. Thus the Indo-west-
Pacific component forms a high proportion of the Solomon Islands fauna. In the
Pacific region, it is interesting that the number of species in common between the

P. E. GIBBS

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THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS in

Solomon Islands and other Island groups lying within the tropics decreases with
increasing distance: the faunas of both New Caledonia and the Marshall Islands
show similar affinities (27-4% and 29-3% respectively) as do the faunas of the
islands of the Central Pacific (15-9%) and Hawaii (15-9%). A relatively high
proportion of the Solomon Islands species extend to Japan (31-2%) but fewer than
expected have been recorded from the east coast of Australia (26-1%) although this
percentage should probably be much higher.

It is difficult to compare these results with those of previous analyses because of
the lack of earlier records from the tropical west Pacific. For example, Knox (1958)
had records of only 81 species from this region (Micronesia - Melanesia) for his analysis
of the distribution of polychaetes in the Indo-Pacific region and of this total 27%
were Indo-Pacific and 31% were cosmopolitan. However the larger number of
records from the present survey indicates that the Indo-Pacific element is proportion-
ally much greater. Interestingly, previous estimates of the Indo-Pacific element
in the faunas of New Zealand and Japan, of 12 % and 30 % respectively (Knox, 1963 ;
Imajima & Hartman, 1964), closely correspond to the overlaps calculated above
between these two regions and the Solomon Islands, the fauna of the latter being
essentially Indo-Pacific.

The records from the Solomon Islands have extended the known distribution of
thirteen species from the Indian Ocean to the West Pacific region. These species
are Lepidasthenia stylolepis, Psammolyce zeylanica, Pareurythoe pitipanaensis,
Phyllodoce fristedti, Syllis longissima, Platynereis insolita, Lumbrineris papillifera,
Marphysa macintoshi, Prionospio ehlersi, Phyllochaetopterus elioti, P. herdmani,
Spiochaetopterus costarum costarum and Megalomma quadrioculatum. A further six
species were known only from Atlantic or Mediterranean waters, namely Trypano-
syllis coeliaca, Sphaerodoridium claparedii, Mastobranchus dollfusi, M. trinchesii,
Abarenicola claparedii, and Megalomma linaresi.

As noted above the fauna of the Solomon Islands appears to be essentially Indo-
Pacific and apart from the new species and subspecies that were discovered, relatively
few of the species remain unrecorded from the Indian Ocean. This Pacific element
in the fauna is represented by such species as Notopygos gregoryi, Eteone japanensis,
Phyllodoce pruvoti, P. (Anaitides) parva, Sigambra hanaokai, Exogone uniformis,
Typosyllis brachycola, T. lucida, Pseudonereis masalacensis, Haploscoloplos bifurcatus,
Pseudopolydora corallicola, Poecilochaetus tropicus, Reteterebella queenslandia,
Megalomma trioculatum and Serpula hartmanae. However most of these species
have few records and probably have yet to be discovered elsewhere.

ECOLOGICAL OBSERVATIONS

Littoral survey

Although it is not practicable to list fully the species taken at each of the littoral
stations, the short accounts that follow are presented as an outline of the ecological
assemblages of species encountered in several of the more important habitats,
in particular, those found on the reef platforms along the more exposed shorelines

H2 P. E. GIBBS

and those composing the infauna of certain sedimentary deposits present in
sheltered coastal areas. The habitats of each species are given in the Systematic
Account. The tidal range in the Solomon Islands is small, being about i-om in
amplitude at springs, and investigations were generally concentrated betwen mid-tide
level and low water mark.

The importance of the boring activities of polychaetes was recognized by Gardiner
(1903) who regarded them as the "prime and most effective agents" in the destruction
of coralline rocks. This view has been supported by Johnson (in Hartman, 1954).
Boring is effected chiefly by the abrasive action of either hard pharyngeal structures,
such as possessed by the nereids and eunicids, or by modified setae, as in certain
spionids and cirratulids (Hartman, 1954). However as pointed out by Utinomi
(1953) it is often difficult to distinguish between those species which actually bore
into the coral from those which utilize and perhaps enlarge the vacated burrows made
by boring species. For example, from its habit the sabellid Hypsicomus phaeotaenia
appears to be a boring species but it does not possess hard chitinized structures that
could be used for penetration. Whether chemical action is employed by such a
species has yet to be investigated. Few polychaetes contribute to reef formation
by virtue of their construction of massed tubes and in the Solomon Islands only
Spirobranchus giganteus appears to be sufficiently common to be considered a minor
contributor to reef growth.

On the reef platforms at moderately or very exposed localities, such as at Matiu,
Batuona and Gizo, sedentary species are surprisingly few although quite extensive
colonies of Phyllochaetopterus socialis are often present in places. On the other hand,
errant species are fairly numerous, particularly the nereids and eunicids, which form
the bulk of the destructive species, including Perinereis nigropunctata, Pseudonereis
anomala, P. masalacensis, P. variegata and perhaps Platynereis insolita, together
with Eunice afra, E. antennata, E. coccinea, E, teniaculata, E. (Palola) siciliensis
and Lysidice collaris. Other species which may contribute to reef breakdown
are Nematonereis unicornis, Arabella mutans, Dodecaceria fistulicola and Hypsicomus
phaeotaenia. Many of these boring species are also common amongst the surface
cover on the reef, that is, amongst algal growth (e.g. Amphiroa), or Phyllochaetopterus
tubes, where they accompany the crevice-dwelling species which utilize such shelter
when the reef is exposed at low-tide. The crevice-dwelling species are many and are
represented by Lepidonotus (Thormora) jukesi, Paleanotus debilis, Eurythoe complanata,
Euphrosine myrtosa, Eulalia (Pterocirrus) magalhaensis, Phyllodoce fristedti, P. pruvoti,
P. quadraticeps , Synelmis albini, Exogone gemmifera, Syllis longissima, Trypanosyllis
coeliaca, Typosyllis alternata, T. armillaris, T. brachycola, T. prolifera, Cirriformia
punctata and Terebella ehrenbergi. Encrusting forms include V ermiliopsis glandigerus
and Hydroides minax. An inconspicuous species of this habitat is Bhawania goodei
which, although found free-living, most frequently occurs as a commensal in the
burrows of the sipunculids Aspidosiphon elegans, Cloeosiphon aspergillum and
Phascolosoma albolineatum.

In sheltered localities the reef platforms are generally covered by a layer of
sediment which is often composed of a wide variety of deposit grades ranging from

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 113

sticky mud to coarse coral debris. At Tetel Island it was possible to investigate an
assortment of deposits in a small area and the following lists of species discovered in
four different grades of sediment give an indication of the diversity of the polychaete
infauna.

(i) In muddy silt-sand at about mid-tide level: iphione muricata, Lepidasthenia
elegans, L. maculata, Sthenelais zeylanica, Eurythoe complanata, Eteone japanensis,
Phyllodoce malmgreni, Langerhansia cornuta, Onuphis (Nothria) holobranchiata,
Malacoceros indicus, Poecilochaetus serpens honiarae n. subsp., P. tropicus,
Mesochaetopterus Sagittarius, Phyllochaetopterus elioti, P. herdmani, Pista dibranchis
n.sp., P. typha and Megalomma vesiculosum. L. elegans and L. maculata were dis-
covered cohabiting the tubes of M . Sagittarius and P. herdmani respectively. Living
under coral boulders lying on this fine deposit, further species were uncovered,
namely, Pseudeurythoe paucibranchiata, Typosyllis alternata, T. brachycola, Perinereis
cultrifera, P. nuntia, Glycera lancadivae, Nematonereis unicornis, Arabella iricolor,
Spiochaetopterus costarum costarum, Hyboscolex longiseta, Dasybranchus caducus,
Lysilla ubianensis and Loimia medusa.

(ii) In coarse coral debris, chiefly Acropora fragments, towards low water mark:
Eurythoe complanata, Notopygos sibogae, Pseudeurythoe paucibranchiata, Leocrates
chinensis, Ceratonereis mirabilis, Eunice aphroditois, E. (Palola) siciliensis, Cirriformia
filigera, Armandia lanceolata, Loima medusa, Reteterebella queenslandia, Branchiomma
cingulata, Hypsicomus phaeotaenia, Megalomma intermedium, M. linaresi, Sabellastarte
sanctijosephi, Spirobranchus giganteus and S. coutierei.

(iii) In shell gravel at low water mark: Lepidonotus (Thormora) jukesi, Amphinome
nigrobranchiata, Pareurythoe pitipanaensis, Leocrates chinensis, Glycera lancadivae,
Eunice (Palola) siciliensis, Onuphis (Nothria) holobranchiata, Mesochaetopterus
sasittarius and Dasybranchus caducus.

(iv) In coarse coral sand just below low water mark: Eurythoe complanata,
Magelona japonica, Poecilochaetus serpens honiarae n. subsp., P. tropicus, Meso-
chaetopterus Sagittarius, Spiochaetopterus costarum costarum, Capitobranchus sp.
Dasybranchus caducus, Mastobranchus trinchesii, PNicomache sp., Pista dibranchis
n. sp. and Megalomma vesiculosum.

At Graham Point in Marau Sound (east Guadalcanal) extensive deposits of silty
sand mixed with shell gravel are colonised by marine angiosperms (Thalassia and
others) and support a rich and varied fauna. Of the polychaetes encountered here
the most interesting are three Lepidasthenia species which were found living as
commensals. L. microlepis and L. stylolepis were both discovered in the burrows
of the sipunculid Siphonosoma vastus while the third species, L. guadalcanalis n.sp.,
was an inhabitant of the burrows of a large enteropneust (probably Balanoglossus
carnosus). The polychaete infauna comprises the following species: Polyodontes
maxillosus, Psammolyce zeylanica, Sthenelais zeylanica, Amphinome nigrobranchiata,
Eurythoe complanata, Pseudeurythoe paucibranchiata, Glycera gigantea, G. lancadivae,
Eunice (Palola} siciliensis, Naineris laevigata, Dispio maroroi n.sp., Scolelepis
squamata mendanai n.subsp., Poecilochaetus tropicus, Mesochaetopterus Sagittarius,
Dasybranchus caducus, Mastobranchus trinchesii, Clymenella (Macroclymene) sp. and

Ii 4 P. E. GIBBS

Pista dibranchis n.sp. Many of these species are also found below coral boulders
together with further species including iphione muricata, Sigalion bandaensis,
Notopygos sibogae, Pareurythoe pitipanaensis, Phyllodoce madeirensis, Notophyllum
splendens, Synelmis albini, Gyptis maraunibinae n.sp., Hesione splendida, Ty posy His
brachycola, Perinereis nigropunctata, Eunice antennata, E. grubei, Lysidice collaris,
Lumbrineris latreilli, Eupolymnia nebulosa, and Euthelepus kinsemboensis. In this
habitat, Hololepidella nigropunctata and H. ophiuricola n.sp. are also found but
living as commensals on ophiuroids.

Dredge Survey of Marovo Lagoon

Along the north-east coast of New Georgia Island and around the north and east
coasts of Vangunu Island in the New Georgia Group, elevated barrier reefs form the

\Tokavai Lagoon
Grassi Lagoon

FIG. 2. Collection localities and positions of the dredge stations in
Marovo Lagoon, New Georgia Group.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 115

seaward edge of a more or less continuous lagoon. This lagoon is about 10 km across
at its widest point, and its depth is less than 50 m over most of its area. Marovo
Lagoon forms the central part of this lagoon, the northern extension being Tokavai
and Grassi Lagoons and the southern part Kolo Lagoon. However, for convenience,
these lagoons will be collectively referred to as Marovo Lagoon. The geomorphology
of the region has been described by Stoddart (1969).

A benthic survey of Marovo Lagoon was carried out in October and November
1965. Qualitative samples of the bottom deposits were taken at 40 stations in
depths from 2 to 35 m, using a small naturalist's dredge. Two stations (ML 218 and
283) were worked in the northern part of the lagoon, the rest being situated in the
central and eastern parts (fig. 2) . The fauna was extracted from the samples (between
10 and 20 1 of sediment) by sieving through a mesh of i-o or 2-0 mm diameter. The
bottom deposits vary from thick terrigenous mud, which occurs close to the main-
land, to coarse coralline sands which are found along the inner margins of the outer
reefs. The station data are given in Table 4.

TABLE 4
Dredge survey of Marovo Lagoon : station data. See fig. 2 for station positions.

Station

Date

Depth

Deposit

(ML)

(1965)

(m)

22.X.

Silty mud

23-x.

Mud

23.X.

Silty mud

25-x.

Mud

25.X.

Mud

25-x.

Mud

26.X.

Silty mud

26.X.

Silty mud

27. x.

Mud

27. x.

Silty mud

27. x.

Silty sand

28.X.

Sand

28.X.

Sand

28.X.

Sand

100

28.X.

Mud

28.X.

Mud

116

29.X.

Silty sand

117

29.X.

Sand

ix8

29.X.

Sand

i34

29.X.

Foram sand

Station

Date

Depth

Deposit

(ML)

(1965)

(m)

155

30.x.

Mud

156

30-x.

Mud

157

3O.x.

Mud

1 88

i.xi

Mud

190

i.xi

Sand

191

i.xi

Silty sand

192

i.xi

Coarse sand

194

2.xi

Mud

195

2.X1

Mud

196

2.xi

Silty sand

203

3-xi

Coarse sand

204

3-xi

Coarse sand

218

8.xi

Mud

228

lo.xi

Mud

229

10. xi

Sand

230

lo.xi

Silty sand

283

1 3-xi

Coarse sand

294

14. xi

Coarse sand

295

14. xi

Coarse sand

296

14. xi

Coarse sand

A total of 65 species were taken during the survey of which 43 were not recorded
from the littoral zone. This category includes all the species records of the families
Nephtyidae (3 species), Lacydoniidae (i), Cossuridae (i), Flabelligeridae (i), Sternas-
pidae (i), Oweniidae (4), Pectinariidae (i), and Ampharetidae (2). In addition the

u6 P. E. GIBBS

survey provided specimens of four new species, namely, Nephtys (Aglaophamus)
munamaorii n.sp., Polydorella novaegeorgiae n.sp., Ophelia koloana n.sp. and
Myriochele heruensis n.sp. and also one new subspecies, Diplocirrus glaucus orientalis.
Of the species taken, ten have not been recorded from the West Pacific, outside of the
Malay Archipelago (Sthenelais heterochela, S. zeylanica, Sthenelanella ehlersi, Chloeia
conspicua, Prionospio steenstrupi malayensis, Armandia longicaudata, Myriochele
eurystoma, Isoldapulchella, Sosane wireni and Lysilla ubianensis) and the distributions
of two species (Lumbrineris papillifera and Prionospio ehlersi} are extended from the
coast of East Africa. A further two species, namely, Pseudopolydora corallicola and
Megalomma trioculatum, have been described only recently from the Marshall Islands.
Haploscoloplos bifurcatus was previously known only from south and south-east
Australia.

The survey provides some indications of the distribution of the commoner species
with respect to the bottom deposits. Arbitrarily defining a "common" species as
one that was present at three or more stations, the following appear to be mainly
restricted to the finer grades of sediments (mud and fine sands) Sthenelais heterochela,
Sthenolepis japonica, Pseudeurythoe paucibranchiata, Sigambra hanaokai, Nephtys
(Aglaophamus) munamaorii n.sp., Paralacydonia weberi, Haploscoloplos bifurcatus,
Prionospio ehlersi, P. pinnata, P. tetelensis n.sp., Diplocirrus glaucus orientalis n.
subsp., Armandia lanceolata, Mastobranchus dollfusi, Isoldapulchella, Loimia medusa,
and Terebellides stroemi. The coarser deposits (medium to coarse sands) appear to
support sparse polychaete populations and only Owenia fusiformis was taken in any
numbers. Species that tolerate a wide range of deposit grades include Glycera
gigantea, G. lancadivae, Eunice marovoi n.sp., Prionospio steenstrupi malayensis,
Armandia leptocirrus, Pectinaria (Pectinaria) antipoda, and Pista dibranchis n.sp.
Apart from these common species, several appear to be locally abundant, to judge
from the large numbers present at single stations. For example, Onuphis (Nothria)
holobranchiata was abundant in the sample of the thick mud at ML 69. Small
species such as Polydorella n'ovaegeorgiae n.sp. and Myriochele heruensis n.sp. were
probably much more abundant than the sample figures at ML 39 and ML 134 suggest,
no doubt many specimens being lost during sieving.

Commensal polychaetes

During the routine sampling of a number of habitats, a total of twelve species,
mainly polynoids, were discovered living in association, apparently as commensals,
with other animals, principally echinoderms and sipunculids. Observations on these
associations have been presented in an earlier paper (Gibbs, 1969) but for the sake
of completeness the commensal species and their hosts are summarized in Table 5.

Although Macnae & Kalk (1962) have described similar associations from the
coast of Mosambique, probably many have yet to be discovered throughout the Indo-
Pacific region. Undoubtedly the echinoderms which act as hosts for G. clavigera,
Hololepidella spp. and P. crinoidicola, are more numerous than the present records
indicate.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 117

TABLE 5

Summary of the commensal polychaetes and their hosts in the Solomon Islands. Abbreviations :
Po - Polychaeta ; Si - Sipunculoidea ; Ga - Gastropoda ; Oph - Ophiuroidea ; Ho - Holothuroidea;

Cr - Crinoidea.
Commensal Host(s)

Bhawania goodei Aspidosiphon elegans (Si)

Cloeosiphon aspergillum (Si)
Phascolosoma albolineatum (Si)

B. pottsiana Eurythoe complanata (Po)

Eunice marovoi n.sp. Cerithium vertagus (Ga)

Gastrolepidia clavigera Bohadschia argus (Ho)

B. graffei (Ho)

Holothuria (Halodeima) atra (Ho)
Stichopus chloronotus (Ho)
Thelenota ananas (Ho)

Hololepidella nigropunctata Ophiarthrum elegans (Oph)

Ophiocoma brevipes (Oph)
O. insularia forma dentata (Oph)
H. ophiuricola n.sp. Macrophiothrix koehleri (Oph)

Ophiarthrum pictum (Oph)

Lepidasthenia elegans Mesochaetopterus Sagittarius (Po)

L. guadalcanalis n.sp. Enteropneust (Balanoglossus carnosus?)

L. maculata Phyllochaetopterus herdmani (Po)

L. microlepis Paraspidosiphon cumingi (Si)

Siphonosoma vastus (Si)

L. stylolepis Siphonosoma vastus (Si)

Paradyte crinoidicola Himerometra robustipinna (Cr)

Recently, Dr R. U. Gooding kindly sent the author three specimens of H. nigropunctata taken from two
A canthaster planci L. collected at Kira-Kira, San Cristobal I. (3.xii.6g) and a further specimen taken from
Diadema savignyi Michelin collected on the Reef Is., Santa Cruz Is. (8.xii.6g).

Brackish water and terrestrial species

Investigations of brackish water were mainly confined to a lagoon at Lunga
Point, near the mouth of the Lunga River on the north coast of Guadalcanal. At
the time of examination (g.ix-n.ix) the salinity of the lagoon was 5'6% , whilst that
of the outside seawater was 34'5% . Three polychaete species were discovered in this
habitat, namely Namalycastis indica, Pseudopolydora sp. and Mercierella enigmatica.
Specimens of N. indica were found burrowing amongst the fibres composing the
outer husk of the Nipa palm nut, many of which were lying waterlogged along the
water's edge. The two other species were present in large numbers within the
interstices of a sponge that was growing around submerged tree roots. The specimens
of Pseudopolydora have proved to represent a new species (W. J. Light, personal
communication) and those of the cosmopolitan M. enigmatica are also of interest
in that they show the characters of Neopomatm and thus support Straughan's (1966)
evidence for the belief that Neopomatus is the warm water form of Mercierella.
Further details of these two species are given below in the Systematic Account.

Wesenberg-Lund (1958) has shown that the majority of freshwater polychaetes

n8 P. E. GIBBS

are nereids and thus it is not surprising that those species which have been discovered
in damp terrestrial habitats also belong to this family, in particular to the subfamily
Namanereinae. In view of the fact that terrestrial forms are well documented from
the East Indies (see Feuerborn, 1932 ; Lieber, 1931 ; Pflugfelder, 1933) their discovery
in the nearby Solomon Islands is to be expected.

Two small specimens belonging to the Namanereinae were separately collected by
members of the Expedition Land Party (Dr. J. Greenslade and Mr. J. Peake) on
Mt. Austen, Guadalcanal and on Kolombangara in the New Georgia Group. Both
were taken from moist leaf litter at an altitude of 350 m. Unfortunately the identity
of these important specimens is uncertain due to damage and although they closely
resemble Namanereis amboinensis (Pflugfelder) they may belong to Cryptonereis
malaitae n.gen., n.sp., described below, which may also be a damp terrestrial form.
Until further specimens from relatively high altitudes are available the status of
these specimens must remain undetermined. Any information concerning their
mode of reproduction would be of great interest.

Spawning of Eunice (Palola) siciliensis

The spawning of the circumtropical species Eunice (Palola} siciliensis is well
documented for the Pacific region and accounts of this phenomenon, which usually
takes place in October or November, have been given by many authors, notably
Burrows (1945, 1955), Gaspers (1961), Gravier (1924), Miller & Pen (1959), Stair (1847)
and Woodworth (1907). Although the Samoan name 'palolo' is the most commonly
used for this species, it has a variety of names, being known as 'paroro' on Rotuma,
'balolo' in the Fiji Islands, and 'hundu' in the New Hebrides. In the Solomon Islands
it has several names ; in the Florida Islands, West Guadalcanal and on Malaita it is
called either 'ogu' or 'odu' and at the eastern end of San Cristobal it is known as
'parenga'. Lever (1945) records that it is called 'orku' on Ulawa Island. Collection
of the 'rising' appears to be a dying custom in the Solomon Islands since many of the
natives interviewed stated that they had not done so for many years and certainly
only the older men could relate the details of the event. From their description of
the rising, it would appear that the spawning of this species appears to be essentially
similar to that described for other areas.

The author could not be present at a rising but fortunately a valuable collection of
the spawning worms was obtained by Wilson Ifunaoa (a schoolboy attached to the
Expedition) from Fanalei Island (also called Falelei or Halelei Island), off the south-
east coast of Maramasike Island, Malaita. This collection was made at the time of
the rising on 2~3.xi.66 and four species are represented - Phyllodoce (Anaitides)
madeirensis, Perinereis cultrifera, Eunice coccinea and Lysidice collaris, in addition to
Eunice (Palola) siciliensis. The details of this material, together with the spawning
times provided by the collector, are given in Table 6. There can be little doubt
that all five species were spawning at the same time because all of the specimens
either contain apparently mature gametes or are in the 'spent' condition. In a

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS

119

similar collection from the New Hebrides, Gravier (1924) records that Lumbrineris
sphaerocephala and Perinereis masalacensis were present.

TABLE 6

Details of the material in the sample of the 'palolo' or 'ogu' rising at Fanalei, Malaita, collected

on 2-3.xi.66 by Wilson Ifunaoa.

Species

Phyllodoce
(Anaitides)
madeirensis

Perinereis
cultrifera

Eunice
coccinea

Eunice

(Palola)

siciliensis

Lysidice
collaris

Material

3 specimens

2 heteronereids

2 posterior
fragments

4 posterior
fragments

i specimen

Time of
spawning (hours)

1830-1900

2OOO-22OO

0000-0600

2OOO-22OO

Condition

Spent

Ripe oocytes
200 (A in diameter

Ripe oocytes
360 (i in diameter

Ripe oocytes
200 (A in diameter

Spent

Local name

Adio

Falisu-ogu

Raka-raka

Falisu-ogu

Also included in the collection of the 'palolo' or 'ogu' is a large specimen of Notopygos
gregoryi, a species hitherto recorded only from Midway Island. It is thought that
this specimen was feeding on the spawning worms as they emerged from the coral.

SYSTEMATIC ACCOUNT

The great majority of the records from the littoral zone were made between mid-
tide level (MTL) and low water mark (LWM). Sublittoral or shallow-water samples
were taken to a maximum depth of 35 m in Marovo Lagoon. For each species the
number of specimens taken at each of the localities or stations (see figs i and 2) is
given : where more than 20 specimens were found the species is indicated as numerous
(num.)

Family APHRODITIDAE

Subfamily POLYNOINAE

Gastrolepidia clavigera Schmarda, 1861

Gastrolepidia clavigera Schmarda, 1861 : 159, pi. 36, fig. 316; Fauvel, 1953 : 51, fig. 22. d-f;
Day, 1967 : 51, fig. 1.5. a-f.

HABITAT. A commensal of holothurians (see Gibbs, 1969 for notes).

izo P. E. GIBBS

RECORDS. On Stichopus chloronotus Brandt - Graham Pt. -8; Maraunibina
Is. - 12; Kalota Is. - 2; on Holothuria (Halodeima) atra Jaeger - Graham Pt. - u ;
Kalota Is. - 5; Auki Hr. - num. ; on Bohadschia argus Jaeger - Maraunibina Is. - 4;
Kalota Is. - 15; on Bohadschia graffei Semper - Maraunibina Is. - 2 ; Yandina - i ;
on Thelenota ananas (Jaeger) - Maraunibina Is. - I.

DISTRIBUTION. Tropical Indo-west-Pacific.

Harmothoe nigricans Horst, 1915

Harmothoe nigricans Horst, 1915 : 14; 1917 : 90, pi. 20, figs 3-4.
HABITAT. Under coral boulders and in beachrock.
RECORDS. Tetel Is. - i ; Komimbo Bay - 4; Lauvie Is. - i.
DISTRIBUTION. East Indies.

Harmothoe sp.

HABITAT. Crevice in reef platform.
RECORD. Maramasike Pg. - i.

NOTES. The specimen has the typical harmothoid prostomium with pronounced
lateral peaks and ventrally inserted lateral tentacles. The elytra number fifteen
pairs and each carries large conspicuous tubercles. The latter are pear-shaped with
wide bases, the largest of them being situated along the posterior margins of the
elytra. Notosetae are numerous and coarsely serrated and neurosetae bidentate
with a fine secondary tooth almost as long as the main one.

The specimen resembles H, impar (Johnston), known from Japan (Imajima and
Hartman, 1964) but differs in that the elytral margins are strongly ciliated.

Hololepidella nigropunctata (Horst, 1915)

Polynoe nigro-punctata Horst, 1915 : 20; 1917 : 104, pi. 21, figs 15-17.

Hololepidella nigropunctata; Day, 1957 : 65, fig. i. a-f; Devaney, 1967 : 287, figs 1-5; Pettibone,

igdga : 50, fig. 2. a-g.

Polyeunoa nigropunctata: Day, 1967 : 54, fig. 1.5. r-u.
Hololepidella minuta: Gibbs, 1969 : 449, fig. 131.

HABITAT. A commensal of ophiuroids living under coral boulders and in crevices
on the reef platform (see Gibbs, 1969).

RECORDS. Graham Pt. - on Ophiocoma brevipes Peters - 8 ; on Ophiocoma insularia
forma dentata Liitken - i ; on Ophiarthrum elegans Peters - 7.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 121

NOTES. In the preliminary report on the commensal species (Gibbs, 1969) these
specimens were referred to Hololepidella minuta (Potts, 1910) because they correspond,
particularly in terms of their setae, with the type specimen of Polynoe minuta in the
British Museum (Natural History). The latter specimen is incomplete however,
having only 24 segments, and thus cannot be referred to the genus Hololepidella
Willey with certainty because the generically characteristic, segmental arrangement
of the elytra (Devaney, 1967; Pettibone, 1969^) cannot be checked. Unfortunately
the type specimens of Polynoe minuta var. oculata Potts (1915) must be considered
lost since they are not present in the British Museum nor in the Zoological Museum
of the University of Cambridge (C. B. Goodhart, pers. comm.). Thus it seems
advisable to refer the Solomon Islands specimens to H. nigropunctata (Horst) and to
regard P. minuta as indeterminable. Possibly further specimens from crinoids at
the type localities of P. minuta (South Male) and P. minuta oculata (Torres Straits)
may clarify the problem.

DISTRIBUTION. Tropical Indo-west-Pacific to Hawaii.

Hololepidella ophiuricola n. sp.

(Fig. 3. A-H)

Hololepidella commensalis: Gibbs, 1969 : 451, fig. 132.

DESCRIPTION. The largest specimen (holotype) measures 15 mm in length, is
2-5 mm wide and has 53 segments bearing 25 pairs of elytra. Smaller specimens
have 41 to 50 segments bearing 19 to 24 pairs of elytra.

The prostomium has marked antero-lateral peaks and two pairs of eyes (fig. 3. A).
The median antenna and tentacular cirri are long and the lateral antennae are short
and ventral in origin. The palps are stout and between 1-5 to 2-0 times the length
of the prostomium. Both dorsal and ventral cirri are relatively long, the former
being about equal to the width of the body, the latter about equal to the length of the
neuropodial lobe.

The elytra are oval in shape and completely cover the body. They have smooth
margins and are transparent except for small, irregular patches of black pigment
contained in polygonal-shaped cells (fig. 3. B, c). As is characteristic of the genus,
the elytra are carried on segments 2, 4, 5, 7, 9, . . . 21, 23, 26, 29, 31, 34, 36 and on
alternate segments to the posterior end. The number of elytra depends on the
segment number and in four of the five specimens there are 24 or 25 pairs on 49 to
53 segments.

The notopodial lobe is rather small (fig. 3. D) and carries 16 to 20 notosetae, each
of which is minutely serrated along one edge (fig. 3. E). The neuropodium is bilobed
with a triangular pre-setal lip and a rounded post-setal lip. There are 3 or 4 supra-
acicular and 6 to 8 sub-acicular neurosetae. Superior neurosetae have about
15 rows of spinules and a smooth, curved tip (fig. 3. F) : inferior neurosetae are shorter
and more slender with minute serrations (fig. 3. G, H).

122 P. E. GIBBS

All specimens have a darkly pigmented dorsal surface the pattern of which may
vary according to the species of the host ophiuroid (see Gibbs, 1969). The cirri and
antennae are dark, as are the palps which also show a white longitudinal line on their
dorsal surface. Apart from a dark circular spot at the base of each parapodium, the
ventral surface is unpigmented.

H. ophiuricola may be distinguished from the type species H. commensalis Willey
and from H. nigropunctata by its notosetae, the ornamentation of which differs in all
three species. Also H. ophiuricola differs from the former species in having a greater
number of notosetae (16 to 20 compared to 8) and from H. nigropunctata in having
unidentate, not bidentate neurosetae. It should be noted that in a recent review of
the genus Hololepidella Pettibone (19690) considers the records of H. commensalis
given by Augener (1922) and Fauvel (1932) from the Indian Ocean are doubtful in
view of the fact that both sets of specimens differ from the type description in having
bidentate neurosetae.

0.5mm

FIG. 3. Hololepidella ophiuricola n.sp. (A) Dorsal view of anterior region showing
pigmentation. (B) Elytron from middle body, (c) Patch of pigment cells on elytron.
(D) Elytrigerous parapodium from middle body. (E) Notoseta. (F-H) Superior and
inferior neurosetae.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 123

HABITAT. A commensal of ophiuroids living under coral boulders and in crevices
on the reef platform.

RECORDS. Graham Pt. - on Macrophiothrix koehleri Clark - 4 (including holo-
type) ; Kalota Is. - on Ophiarthrum pictum Miiller and Troschel - i.
British Museum (Natural History) Registration No. Holotype 1970-20

Paratypes 1970-21-24

Iphione muricata (Savigny, 1818)

Iphione muricata: Fauvel, 1953 : 3 2 > fig- X 3- a ~ e ; Day, 1967 : 43, fig. 1.3. a-f.
HABITAT. Under coral boulders in muddy silt and sand.

RECORDS. Tetel Is. - 3; Komimbo Bay - 3; Maraunibina Is. - 2; Graham
Pt. - 24.

DISTRIBUTION. Tropical Indo-west-Pacific.

Lepidasthenia elegans (Grube, 1840)

Lepidasthenia elegans : Potts, 1910 : 342, pi. 19, fig. 16, pi. 20, fig. 32; Fauvel, 1923 : 88, fig. 33.
a-g; Day, 1967 : 90, fig. 1.16. i-m.

HABITAT. In silt and cohabiting a tube of Mesochaetopterus Sagittarius at LWM.

RECORDS. Tetel Is. - 2.

DISTRIBUTION. Mediterranean; Indo-west-Pacific.

Lepidasthenia guadalcanalis n. sp.

(Fig. 4. A-F)

Lepidasthenia mossambica: Gibbs, 1969 : 453, fig. 134.

DESCRIPTION. The holotype is the largest of the six specimens and measures 85 mm
for about 125 segments. Preserved in alcohol the colour is pale brown with darker
bars across the anterior segments. The fourth pair of elytra is coloured reddish-
brown (fig. 4. A) but the other elytra are colourless.

The prostomium is roughly hexagonal in shape and has two pairs of eyes situated
on the antero-lateral and posterior margins (fig. 4. B). The median and lateral
antennae are terminal in origin and are about one half the length of the stout palps.
The tentacular cirri are slightly longer than the antennae. The posterior margin
of the prostomium is hidden by a conspicuous occipital flap which is heavily papil-

I2 4

P. E. GIBBS

lated. Similar papillae are to be found along the anterior and posterior edges of the
anterior segments on their dorsal side and a few are present on the dorsal and ventral
surfaces of the neuropodial lobes.

The elytra, which are arranged segmentally in the typical polynoid sequence, are
thin, circular in shape and do not cover the mid-dorsal region of the body. They
have smooth margins and lack papillae. Only the fourth pair, on segment 7, is
pigmented and the others are almost transparent.

The dorsal cirri are about equal in length to the neuropodial lobes. The noto-
podial lobes are small and supported by an aciculum but the neuropodial lobes are
well-developed with rounded, subequal, pre-setal and post-setal lips and short
ventral cirri (fig. 4. c).

Notosetae appear to be entirely lacking. In the anterior segments the neurosetae
are slender with a small subterminal tooth (fig. 4. D, E) and number about 30 per
neuropodium. However in the middle body segments, the neurosetae are much
stouter, fewer in number (14 or 15 per neuropodium) and lack subterminal teeth
(fig. 4. F). The number of rows of spinules possessed by these setae is similar

0.5mm

FIG. 4. Lepidasthenia guadalcanalis n.sp. (A) Dorsal view of anterior region showing
pigmentation of fourth pair of elytra. (B) Head region, (c) Parapodium from middle
body. (D-E) Superior neuroseta from anterior segment. (F) Superior neuroseta from
middle body segment.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 125

throughout the body, the superior ones having about 7, the inferior ones 4 or 5.
Neither slender superior neurosetae in the anterior segments nor giant neurosetae
in the posterior region could be found.

In possessing a distinctly papillated occipital flap, L. guadalcanalis resembles three
other species, namely, L. michaelseni Augener, L. terrareginae Monro and L. mossambica
Day. It differs from the former two species in lacking slender superior neurosetae
and in having unidentate neurosetae in the middle segments. L. guadalcanalis is
close to L. mossambica, to which these specimens were provisionally assigned (Gibbs,
1969), but a comparison of the type specimen of L. mossambica (in the British
Museum, Natural History) has revealed important differences in the structure of the
setae. In the middle segments of L. guadalcanalis the neurosetae are much stouter,
fewer in number and have fewer rows of spinules (7 compared to 10 or 12) than in
L. mossambica and the presence of small subterminal teeth on the neurosetae of the
anterior segments of L. guadalcanalis also separates the two species. Furthermore
the distinctive coloration of the fourth pair of elytra appears to be a characteristic
feature of L. guadalcanalis.

HABITAT. A commensal of a large enteropneust (Balanoglossus carnosus?} living
in silty sand (Thalassia flat).

RECORDS. Graham Pt. - 4; Fintry Pt. - 2 (including holotype).
British Museum (Natural History) Registration No. Holotype 1970-25

Paratypes 1970-26-28

Lepidasthenia maculata Potts, 1910

Lepidasthenia maculata Potts, 1910 : 344, pi. 20, fig. 33, pi. 21, fig. 51; Fauvel, 1953 : 58, fig. 27.
h-k; Day, 1967 : 92, fig. 1.16. s-v.

HABITAT. In coarse sand, silt and within a tube of Phyllochaetopterus herdmani.

RECORDS. Haroro - i ; Tetel Is. - 2.

DISTRIBUTION. North Atlantic; Indian Ocean; Indo-China.

Lepidasthenia microlepis Potts, 1910

Lepidasthenia microlepis Potts, 1910 : 343, pi. 19, fig. 17, pi. 21, fig. 52; Fauvel, 1953 : 57, fig. 26.
e-f; Day, 1967 : 90, 1.16. e-h.

HABITAT. A commensal of sipunculids. Hosts include Siphonosoma vastus
(Selenka & Biilow) living in silty sand (Thalassia flat) and Paraspidosiphon cumingi
(Baird) boring in a Porites boulder.

RECORDS. Graham Pt. - 4.
DISTRIBUTION. Tropical Indo-west-Pacific.

126 P. E. GIBBS

Lepidasthenia stylolepis Willey, 1907

Lepidasthenia stylolepis Willey, in Lloyd, 1907 : 260, figs 1-4; Gibbs, 1969 : 453, fig. 134.

HABITAT. A commensal of the sipunculid Siphonosoma vastus, living in silty sand
(3 out of 7 specimens were discovered in association; hosts were not found for the
other 4 specimens).

RECORDS. Graham Pt. - 7.

NOTES. The similarity of Perolepis regularis Ehlers and Lepidasthenia sibogae
Horst to L. stylolepis has been referred to in an earlier paper (Gibbs, 1969).

DISTRIBUTION. Persian Gulf; PEast Africa; PEast Indies.

Lepidonotus (Thormora) jukesi (Baird, 1865)

Lepidonotus (Thormora) jukesi: Fauvel, 1953 : 37, fig. 13. o-r; Day, 1967 : 80, fig. 1.13. g-m.

HABITAT. Under coral boulders, in crevices within reef platform, amongst
Phyllochaetopterus socialis tubes and algal cover (Amphiroa) from MTL to LWM.

RECORDS. Tetel Is. - i ; Matiu Is. - 13 ; Batuona Is. - 17 ; New Manra - 5 ;
Komimbo Bay - i ; Mamara Pt. - I ; Lingatu - 2 ; Maramasike Pg. - 3.

DISTRIBUTION. Indo-west-Pacific.

Paradyte crinoidicola (Potts, 1910)

Polynoe crinoidicola Potts, 1910 : 337, pi. 18, fig. 10, pi. 20, fig. 30, pi. 21, figs 39-41.
Scalisetosus longicirrus : Fauvel, 1953 : 50, fig. 22. a-c; Day, 1967 : 58, fig. 1.7. a-f.
Paradyte crinoidicola: Pettibone, 1969 : 13, fig. 7. a-g.

HABITAT. A commensal of crinoids; on Himerometra robustipinna (P. H.
Carpenter).

RECORDS. Maraunibina Is. - 5 specimens from two hosts.

NOTES. Following Fauvel (1953) and Day (1967) P. crinoidicola was previously
referred (Gibbs, 1969) to S. longicirrus (Schmarda) which Pettibone (1969) considers
indeterminable.

DISTRIBUTION. Indo-west-Pacific.

Paralepidonotus ampulliferus (Grube, 1878)

Polynoe ampullifera Grube, 1878 : 35, pi. 3, fig. 5.

Harmothoe ampullifera: Fauvel, 1953 : 43, fig. i8.d; Pillai, 1965 : 117, fig. 3. d-g, fig. 4. a-b.

Paralepidonotus ampulliferus: Horst, 1917 : 76; Day, 1967 : 47, fig. 1.4. a-f.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 127

HABITAT. Undersurface of coral boulder on reef platform.

RECORD. Kalota Is. - i.

DISTRIBUTION. Tropical Indo-west-Pacific.

Paralepidonotus indicus (Potts, 1910)

Lagisca indica Potts, 1910 : 338, pi. 19, fig. 13, pi. 21, figs 46-47.
Paralepidonotus indicus: Day, 1967 : 48, fig. 1.4. g-k.

HABITAT. Silty sand with coral debris at LWM.
RECORD. Komimbo Bay- i.
DISTRIBUTION. Mogambique; Maldive Is.

Subfamily POLYODONTIDAE
Polyodontes maxillosus (Ranzani, 1817)

Polyodontes maxillosus: Fauvel, 1923 : 97, fig. 37. a-n; 1953 : 71, fig. 32. a-n.

HABITAT. Silty sand with shell and coral fragments at LWM; coarse sand at
1 8 m depth.

RECORDS. Graham Pt. - i ; Fintry Pt. - i ; ML 204 - i.

NOTES. Fauvel records that this species may grow to i m in length. The largest
of the three specimens from the Solomon Islands measures only 12 cm: its felt-like
tube had a diameter i-o to 1-5 cm and extended to a depth of about 30 cm into the
deposit at Graham Pt.

DISTRIBUTION. North Atlantic ; Mediterranean ; Indian Ocean ; Indo-China.

Polyodontes melanonotus (Grube, 1876)

Polyodontes melanonotus: Fauvel, 1953 : 72, fig. 33. c-g; Day, 1967 : 96, fig. 1.17. g-n.

HABITAT. Mud at n m; coarse sand at 18 m.
RECORDS. ML 194 - i ; ML 204 - i.

NOTES. Specimen from ML 194 is 3-5 cm long: it was removed from a tube
about 15 cm in length, i-o cm in diameter, composed of consolidated mud.

DISTRIBUTION. West Africa; Indo-west-Pacific.

128 P. E. GIBBS

Subfamily SIGALIONINAE
Psammolyce zeylanica Willey, 1905

Psammolyce zeylanica Willey, 1905 : 255, pis 1-2, figs 33-43; Fauvel, 1953 : 68, fig 31.!.
HABITAT. Silty sand (Thalassia flat).
RECORDS. Graham Pt. - 2.
DISTRIBUTION. Ceylon.

Sigalion bandaensis Horst, 1917

Sigalion bandaensis Horst, 1917 : no, pi. 22, figs 4-5.

HABITAT. Under boulder on silty sand with coral debris at LWM.

RECORD. Graham Pt. - i.

DISTRIBUTION. East Indies; East Australia.

Sthenelais heterochela Horst, 1917

Sthenelais heterochela Horst, 1917 : 113, pi. 23, figs 3-6.

HABITAT. Mud and silty sand, 2-11 m depth.
RECORDS. ML 191 - i ; ML 194 - i ; ML 195 - i.
DISTRIBUTION. East Indies.

Sthenelais zeylanica Willey, 1905

Sthenelais zeylanica Willey, 1905 : 258, pi. 2, fig. 48; Fauvel, 1953 : 62, fig. 29. a.
HABITAT. Silty sand, MTL to LWM ; sand, 5-16 m depth.

RECORDS. Tetel Is. -i; Komimbo Bay -4; Graham Pt. -2; Fintry Pt. -3;
ML 134 - 2 ; ML 190 - i.

NOTE. In addition to the two slender stylodes arising from the base of the ventral
cirrus, these specimens have a third appendage with a bulb-shaped ending.

DISTRIBUTION. India; Palau Islands.

Sthenelanella ehlersi (Horst, 1916)

Euleanira ehlersi Horst, 1916 : 12; 1917 : 122, pi. 27, figs 1-5; Day, 1967 : 101.
Sthenelanella ehlersi: Pettibone, 19696 : 434, figs 4-5.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 129

HABITAT. Mud and silty sand, 18-24 m depth.
RECORDS. ML 188 - 3; ML 230- i.
DISTRIBUTION. Natal; East Indies.

Sthenolepis japonica (Mclntosh, 1885)

Leanira japonica Mclntosh, 1885 : 154, pi. 22, fig. 3, pi. I4A, figs 1-2; Fauvel, 1953 : 69, fig.

33-a-b.
Sthenolepis japonica: Imajima & Hartman, 1964 : 43; Day, 1967 : 112.

HABITAT. A characteristic species of fine deposits in the sublittoral zone, thick
mud to silty sand, 2-33 m depth.

RECORDS. ML 56 - 2 ; ML 69 - i ; ML 100 - 5 ; ML 155 - 2 ; ML 156 - 2 ; ML 157 -
i; ML 188-2; ML 190 -2; ML 191-1; ML 195 -3; ML 196 -3; ML 218 -4;
ML 228 - i.

DISTRIBUTION. Indo-west-Pacific to Japan.

Thalenessa digitata Mclntosh, 1885

Thalenessa digitata Mclntosh, 1885 : 140, pi. 22, fig. 2, pi. 23, figs 5-7, pi. 25, figs 4-5, pi. I3A,
figs 7-10; Willey, 1905 : 260, pi. 2, figs 50-52; Imajima & Hartman, 1964 : 46.

HABITAT. Coarse sand, 2-4 m depth.
RECORDS. ML 203 - 2 ; ML 296 - i.
DISTRIBUTION. Ceylon ; Admiralty Is. ; Japan.

Family PALMYRIDAE
Bhawania goodei Webster, 1884

Bhawania cryptocephala Gravier, 1901 : 263, pi. 10, figs 152-156, text-figs 280-285; Fauvel,

1953 : 79, ng. 36. e-i.
Bhawania goodei: Day, 1953 : 407 (synonymy); 1967 : 118, fig. 2.1. a-f.

HABITAT. Cohabiting burrows of coral-boring sipunculids; hosts include Aspido-
siphon elegans (Chamisso & Eysenhardt), Cloeosiphon aspergillum (Quatrefages) and
Phascolosoma albolineatum (Baird). Also found free-living in crevices and amongst
tubes of Phyllochaetopterus socialis.

RECORDS. Matiu Is. - 30; New Manra - 5 ; Mamara Pt. - i ; Maramasike Pg. - 2.

DISTRIBUTION. Circumtropical.

130 P. E. GIBBS

Bhawania pottsiana Horst, 1917

Bhawania cryptocephala: Potts, 1910 : 328.

Bhawania cryptocephala var. pottsiana Horst, 1917 : 137.

Bhawania sp.: Gibbs, 1969 : 454.

HABITAT. Sedimented crevices in Porites boulders; commensal (?) with Eurythoe
complanata (see Gibbs, 1969).

RECORDS. Graham Pt. - 2.

NOTES. The larger of the two specimens measures 45 mm in length. Both
possess very slender setae with hair-like appendices in the inferior part of the
neuropodium, which are not present in B. goodei. This additional type of seta was
first noticed by Potts (1910) in a specimen from Zanzibar which he attributed to
B. cryptocephala. However Horst (1917), in recording a similar specimen from the
Celebes Is., gave this form varietal status, naming it B. cryptocephala var. pottsiana.
Day (1953) has shown B. cryptocephala to be a synonym of B. goodei, but since the
variety pottsiana is quite distinct from its stem species, its subspecific status is here
raised to specific rank.

DISTRIBUTION. Zanzibar; East Indies.

Paleanotus debilis (Grube, 1855)
Paleanotus debilis: Day, 1962 : 635 (synonymy); 1967 : 117, fig. 2.1. g-k.

HABITAT. Amongst Phyllochaetoptems socialis tubes at LWM; on Halichondria
sp. at 5 m depth.

RECORDS. Matiu Is. - i ; Batuona Is. - i ; Yandina - 2.
DISTRIBUTION. North Atlantic; Mediterranean; Indo-Pacific.

Family AMPHINOMIDAE
Amphinome nigrobranchiata Horst, 1912

Amphinome nigrobranchiata Horst, 1912 : 39, pi. 10, figs 17-20.
HABITAT. Under coral boulders; in shell gravel deposits.
RECORDS. Tetel Is. - 2 ; Graham Pt. - 2 ; Maraunibina Is. - i.
DISTRIBUTION. East Indies.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 131

Chloeia conspicua Horst, 1910

Chloeia conspicua Horst, 1910 : 173; 1912 : 20, pi. 7, fig. 5, pi. 8, figs 4-5.

HABITAT. Silty sand at 2 m depth.
RECORD. ML 196 - i.
DISTRIBUTION. East Indies.

Euphrosine foliosa Audouin & Milne-Edwards, 1833

Euphros ine foliosa : Fauvel, 1923 : 136, fig. 49. a-g; 1953 : 102, fig. 48. a-h.
HABITAT. Under coral boulder at LWM.
RECORD. Maraunibina Is. - i.
DISTRIBUTION. Atlantic Ocean; Mediterranean; Indo-west-Pacific.

Euphrosine myrtosa Savigny, 1818

Euphrosine myrtosa : Fauvel, 1923 : 139, fig. 49. k-n; 1953 : 101, fig. 48. k-n; Day, 1967 : 127,
fig. 3.1.2.

HABITAT. Amongst Phyllochaetopterus socialis tubes; sedimented crevice in
Porites boulder.

RECORDS. Batuona Is. - i ; Graham Pt. - i.

DISTRIBUTION. West Africa; Mediterranean; Indo-west-Pacific.

Eurythoe complanata (Pallas, 1766)
Eurythoe complanata : Fauvel, 1953 : 83, fig. 38. b-m; Day, 1967 : 128, fig. 3.2. a-h.

HABITAT. A wide variety of situations, including under coral boulders, in crevices
in reef platform and in beachrock, in Acropora rubble and amongst Phyllochaetopterus
socialis tubes and Amphiroa.

RECORDS. Tetel Is. -10; Komimbo Bay -20; Matiu Is. -num. ; Batuona Is. -
num. ; New Manra - i ; Graham Pt. - 6; Maraunibina Is. - 5 ; Lauvie Is. - i.

DISTRIBUTION. Circumtropical.

? Eurythoe sp.

HABITAT. Under coral boulders in silty sand.
RECORDS. Maraunibina Is. - 4.

NOTES. The specimens measure between 10 and 20 mm long and about 2 mm
wide and have 57 to 67 segments. A sinuous caruncle extends through setiger 2

132 P. E. GIBBS

and the branchiae commence between setigers 9 and 13, extending to the posterior
end. Notosetae are of three types, namely (i) stout spines, (ii) harpoon setae and
(iii) long, fine, smooth capillaries. Neurosetae are of two types - (i) furcate, with a
short secondary spur and (ii) long, fine capillaries with faintly serrated blades, some
with a stout spur.

The generic identity of these specimens is in question since in the genus Eurythoe
the branchiae commence on setigers 1-3 (Day, 1967). It is possible that these
relatively small specimens are juveniles and because of this, the erection of a new
genus must be postponed until further material is available.

Notopygos gregoryi Holly, 1939

Notopygos gregoryi Holly, 1939 : 265, fig. i. a-f.

HABITAT. Not known.
RECORD. Fanalei - 1.

NOTES. The specimen of this elegant species was collected during the 'palolo'
rising at Fanalei between 2000 and 2200 hours on 2.xi.66. It is 160 mm long and
25 mm across the body. All details correspond to the description of the holotype.

DISTRIBUTION. Midway Is. (one specimen).

Notopygos sibogae Horst, 1911

Notopygos sibogae Horst, 1911 : 245; 1912 : 27, pi. 9, figs 4-5.

HABITAT. In Acropora rubble ; within sedimented crevices on the undersurfaces
of Porites boulders.

RECORDS. Tetel Is. - i ;, Graham Pt. - 3.

NOTES. Hartman (1966) comments that the differences in the specific characters
in Notopygos i.e. number of segments and the position of the anal pore, may reflect
differences in the growth stages. However in all four specimens in the collection of
N. sibogae the anal pore is situated on the anterior margin of setiger 23 although the
lengths vary from 10 to 20 mm.

DISTRIBUTION. East Indies.

Pareurythoe pitipanaensis De Silva, 1965

Pareurythoe pitipanaensis De Silva, 1965 : 540, fig. 3. a-k.

HABITAT. Under coral boulders, in shell gravel and within the interstices of the
coral Galaxea.

RECORDS. Tetel Is. - i ; Graham Pt. - 5 ; Maraunibina Is. - 3.
DISTRIBUTION. Ceylon.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 133

Pseudeurythoe paucibranchiata Fauvel, 1932

Pseudeurythoe paucibranchiata Fauvel, 1932 : 47, pi. i, figs 3-4, text-fig. 8. a-e; 1953 : 86, fig.
39. a-b, fig. 40. a-e.

HABITAT. In silty deposits, particularly under coral boulders; in Acropora
rubble ; in muds and fine sands, 2-26 m depth.

RECORDS. Tetel Is. - 3 ; Komimbo Bay - 12 ; Graham Pt. - 7 ; Maraunibina Is. -
4; ML 37-2; ML 41 - i; ML 68- i; ML 96-1; ML no- 15; ML 134- i; ML 156-
3 ; ML 157 - 6 ; ML 194 - i ; ML 195 - i ; ML 218 - i ; ML 228 - 4.

NOTES. Most of the specimens show a button-like caruncle set into the first setiger
but some are so contracted that this feature is not visible. Small specimens about
5 mm in length have 4 or 5 pairs of branchiae, whilst in medium-sized specimens up
to 10 mm long, the branchiae continue to setiger 16 or 17. The largest specimens
come from the littoral zone (Marau Sound) ; these measure 35 mm long and have
branchiae up to setiger 27, as typical for P. paucibranchiata.

There is some doubt as to whether all of the specimens in this series should be
referred to P. paucibranchiata since the position of the prostomium in relation to the
first segments and the presence of a caruncle, two major specific characters, are
difficult to determine in strongly contracted specimens. Further, the number of
branchiae varies greatly with size and cannot be regarded as specific, particularly in
small specimens. The setae also are not diagnostic, according to Wesenberg-Lund
(1949). On account of these considerations, it seems probable that a number of
Pseudeurythoe species are synonyms of P. oculifera (Augener) , the description of which
was based on a small, probably juvenile, specimen.

DISTRIBUTION. Indian Ocean ; Indo-China.

Family PHYLLODOGIDAE
Eteone japanensis Mclntosh, 1901

Eteone japanensis Mclntosh, 1901 : 222, pi. i, fig. 2; Imajima & Hartman, 1964 : 60.,

HABITAT. Silty sand.
RECORD. Tetel Is. - i.

NOTES. The specimen agrees with those details given in the type description and,
in addition, possesses a smooth proboscis. However, since the characters of the
proboscis in this species are unknown the identification must be regarded as a
provisional one.

DISTRIBUTION. Japan Sea.

134 p - E - GIBBS

Eulalia albopicta Marenzeller, 1879

Eulalia albopicta Marenzeller, 1879 : 128, pi. 3, fig. 3; Fauvel, 1953 : 123, fig. 60. a-b.
HABITAT. Cohabiting a Phyllochaetopterus elioti tube from silty sand.
RECORD. Fintry Pt. - i.
DISTRIBUTION. Indian Ocean; Indo-China; Japan.

Eulalia viridis (Linnaeus, 1767)

Eulalia viridis : Fauvel, 1923 : 160, fig. 57. a-h; 1953 : 122, fig. 6.1. a-h; Imajima & Hartman,
1964 : 63.

HABITAT. Crevice in reef platform.
RECORD. Maramasike Pg. - i.
DISTRIBUTION. Cosmopolitan.

Eulalia (Pterocirrus) magalhaensis Kinberg, 1866

Eulalia (Pterocirrus) magalhaensis : Kinberg 1858-1910 : 55, pi. 33, fig. i; Fauvel, 1953 : 124,
fig. 62. a-h.

HABITAT. Amongst Phyllochaetopterus socialis tubes and algal growth (Amphiroa)
towards LWM.

RECORDS. Cape Esperance - i ; Matiu Is. - 2; Batuona Is. - i ; New Manra - i.

NOTE. All five specimens are small-sized (3 to 7 mm) but conform to the species
description.

DISTRIBUTION. Indo-Pacific.

Notophyllum splendens (Schmarda, 1861)

Macrophyllum splendens Schmarda 1861 : 82, pi. 29, fig. 227.

Notophyllum splendens: Fauvel, 1953 : 126, fig. 60. c; Day, 1967 : 151, fig. 5.3. k-n.

HABITAT. Under coral boulder; interstices of the coral Galaxea.
RECORDS. Graham Pt. - 2.
DISTRIBUTION. Indo-west-Pacific.

Phyllodoce fristedti Bergstrom, 1914

Phyllodoce fristedti : Fauvel, 1953 : 118, fig. 58. a-b; Day, 1967 : 147, fig. 5.2. k-m.

HABITAT. Crevices in reef platform and beachrock.
RECORDS. Matiu Is. - i ; Lauvie Is. - i.
DISTRIBUTION. Indian Ocean.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 135

Phyllodoce malmgreni Gravier, 1900

Phyllodoce malmgreni Gravier, 1900 : 207, pi. 10, figs 2931, text-figs 66-69; Fauvel, 1953 : JI 7
fig. 56. h; Day, 1967 : 147, fig. 5.2. n-p.

HABITAT. Silty sand at MTL; mud at 2 m depth.
RECORDS. Tetel Is. - 2 ; ML 68 - i.
DISTRIBUTION. Indian Ocean; East Australia.

Phyllodoce pruvoti Fauvel, 1930
Phyllodoce pruvoti Fauvel 1930 : 512, fig. i. a-f, fig. 2. a-c; 1947 : 28, fig. 23.

HABITAT. Crevices in reef platform ; amongst Phyllochaetopterus socialis tubes.
RECORDS. Matiu Is. - i ; Batuona Is. - i ; New Manra - i.
DISTRIBUTION. Tropical West Pacific.

Phyllodoce quadraticeps Grube, 1878

Phyllodoce quadraticeps Grube, 1878 : 98, pi. 6, fig. 2; Fauvel, 1953 : 116, fig. 56. f-j; Day, 1967 :
145, fig. 5.2. h-j.

HABITAT. Crevices in reef platform and in beachrock, MTL to LWM.
RECORDS. Kokomtambu Is. - i ; Matiu Is. - 10; Lauvie Is. - i.

NOTE. This species is often found crawling over the surface of exposed coral
during the low-tide period.

DISTRIBUTION. Tropical Indo-west-Pacific.

Phyllodoce (Anaitides) madeirensis Langerhans, 1880

Phyllodoce (Anaitides) madeirensis: Fauvel, 1953 : 120, fig. 59. d-h; Day, 1967 : 145, fig. 5.2. d-g.

HABITAT. In Acropora rubble, under coral boulders and in beachrock crevices.
RECORDS. Komimbo Bay - 2 ; Lauvie Is. - 2 ; Graham Pt. - 2 ; Fanalei - 3.
NOTES. The Fanalei specimens were collected between 1830 and 1900 hours
during the 'palolo' rising on 2.xi.66 and are 'spent'. Local name is 'adio'.

DISTRIBUTION. Cosmopolitan.

136 P. E. GIBBS

Phyllodoce (Anaitides) parva (Hartmann - Schroder, 1965)
Anaitides parva Hartmann - Schroder, 1965 : 88, figs 7-9; Hartman, 1966 : 183.

HABITAT. Medium sand at 5 m.
RECORD. ML 190-1.
DISTRIBUTION. Hawaii.

Phyllodoce (Genetyllis) castanea (Marenzeller, 1879)

Carobia castanea Marenzeller, 1879 : 127, pi. 3, fig. 2; Izuka, 1912 : 199, pi. 21, fig. 3.
Phyllodoce (Genetyllis) castanea: Fauvel, 1953 : 115, fig. 56. a-c; Day, 1967 : 149, fig. 53. d-f.

HABITAT. Sand at n m.
RECORD. ML 98-1.
DISTRIBUTION. Indo-west-Pacific.

Phyllodoce (Genetyllis) gracilis Kinberg, 1866

Phyllodoce gracilis : Kinberg, 1858-1910 : 55, pi. 22, fig. 3; Monro, 1939 : 173, fig. 3. a-c; Fauvel,
1953 : 117. fi g- 57- a-g.

HABITAT. Crevice in reef platform.
RECORD. Lingatu - 1.
DISTRIBUTION. Tropical Indo-Pacinc.

Family PILARGIDAE
Sigambra hanaokai (Kitamori, 1960)

Ancistrosyllis hanaokai Kitamori, i96oa : 1086, fig. i. a-h.
Sigambra hanaokai: Pettibone, 1966 : 181 (synonymy).

HABITAT. Mud, 2-20 m depth.

RECORDS. ML 39 - 3 ; ML 40 - i ; ML 41 - 10 ; ML 68 - 5 ; ML 157-1.

NOTES. Larger specimens - up to 14 mm in length - were found inhabiting thin

membraneous tubes covered in mud particles.

DISTRIBUTION. Japan.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 137

Synelmis albini (Langerhans, 1881)

Ancistrosyllis rigida Fauvel, 1919 : 337, fig. i. a-e; 1953 : no, fig. 53. a-e; Day, 1967 : 215.
Synelmis albini: Pettibone, 1966 : 191, figs 19-21 (synonymy).

HABITAT. Crevices in reef platform; amongst Phyllochaetopterus socialis tubes;
in shell gravel and coarse sand.

RECORDS. Komimbo Bay - 2 ; Matiu Is. - i ; Batuona Is. - 4; Mamara Pt. - I ;
Graham Pt. - i ; Maramasike Pg. - i.

DISTRIBUTION. Circumtropical.

Family HESIONIDAE
Gyptis capensis (Day, 1963)

Oxydromus capensis Day, 1963 : 397, fig. 4. e-j.
Gyptis capensis: Day, 1967 : 231, fig. 11.2. l-o.

HABITAT. Under coral boulders on silty sand, MTL to LWM.
RECORDS. Maraunibina Is. - 3.

NOTES. A complete specimen measures 15 mm in length for about 70 segments.
Notosetae are present from setiger 5.

DISTRIBUTION. South Africa.

Gyptis maraunibinae n. sp.

(Fig. 5. A-F)

DISTRIBUTION. The holotype is 45 mm long for 150 segments and the body is
about 3 mm wide.

The prostomium is rectangular, its width being about 1-5 times the length (fig. 5. A).
Three antennae arise from the anterior margin of the prostomium; the laterals
are about twice the length of the median. The two biarticulate palps, each com-
posed of a short palpostyle and longer palpophore, are stouter and slightly longer
than the lateral antennae. There are two pairs of eyes and those of the anterior
pair are reniform in shape and are larger than those of the posterior pair. The
proboscis carries ten papillae, each of which is broad and flap-like (fig. 5. B). Jaws
are lacking, but there is a horny rim around the ventral and ventro-lateral margins
of the proboscis.

The tentacular cirri number eight pairs and each is faintly but closely annulated.
The dorsal pair of the second segment is the longest, reaching back to about
setiger 15.

138 P. E. GIBBS

The first 4 or 5 setigers are uniramous, becoming biramous when a notopodial
papilla appears on the antero-ventral side of the cirrophore from about setiger 6.
The notopodia are small throughout the length of the body but the neuropodia are
well developed, each with a pointed pre-setal lip and a rounded post-setal lip (fig. 5. c).
Ventral cirri are about equal to the neuropodial lobe in length. In the middle
of the body, notosetae include 2 or 3 rather stout, smooth spines (fig. 5. E) and 4 or
5 forked setae (fig. 5. D) ; neurosetae are falcigerous, each with a minutely serrated
blade of varying length and with a bidentate tip consisting of a strong terminal
tooth and a slender secondary one (fig. 5. F).

The large, lobular, papillae on the proboscis are sufficient to distinguish G.
maraunibinae from other Gyptis species. In other details the species resembles G.
capensis (Day).

HABITAT. Silty sand with shell gravel below coral boulder on reef platform.

RECORD. Graham Pt. - i.

British Museum (Natural History) Registration No. Holotype 1970-29

1.0mm

0.5mm

=1
o

0.5mm

FIG. 5. Gyptis maraunibinae n.sp. (A) Dorsal view of prostomium. (B) Anterior view of
proboscis, (c) Parapodium from middle body. (D-E) Notopodial forked seta and spine.
(F) Neuropodial falciger.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 139

Hesione splendida Savigny, 1818

H esione pantherina : Fauvel, 1953 : IO 4' fig- 49- a "g-
Hesione splendida: Day, 1967 : 228, fig. 11.2. a-c.

HABITAT. In Acropora rubble and shell gravel; in crevices and under coral
boulders.

RECORDS. Honiara - i ; Komimbo Bay - i ; Graham Pt. - 14.
DISTRIBUTION. North Atlantic ; Mediterranean ; Indo-west-Pacific.

Leocrates chinensis Kinberg, 1866

Leocrates claparedii: Fauvel, 1953 : 106, fig. 50. c-g; Day, 1967 : 230, fig. 11.2. g-k.
Leocrates chinensis: Imajima & Hartman, 1964 : 82 (synonymy).

HABITAT. In shell gravel and Acropora rubble; under coral boulders.

RECORDS. Kokomtambu Is. - i ; Tetel Is. - 4; Komimbo Bay - i ; Fintry Pt. -
i ; Maramasike Pg. - i.

DISTRIBUTION. West Africa; Mediterranean; tropical Indo-west-Pacific.

Family SYLLIDAE

The species listed below for this family were identified by Dr. Minoru Imajima.
A detailed account of this material will be published by Dr. Imajima at a later date.

Subfamily AUTOLYTINAE
Autolytus sp.

HABITAT. On sponge Halichondria sp. at 5 m depth.
RECORD. Yandina- i (damaged).

Subfamily EXOGONINAE
Brania clavata (Claparede, 1863)

Grubea clavata: Fauvel, 1923 : 296, fig. 114 a-e.
Brania clavata: Imajima, 1966 : 393, fig. i a-g.

HABITAT. Coarse sand at LWM.

RECORD. Komimbo Bay - i.

DISTRIBUTION. North Atlantic; Mediterranean; Japan.

i 4 o P. E. GIBBS

Exogone gemmifera Pagenstecher, 1862

Exogone gemmifera : Fauvel, 1923 : 305, fig. 117. a-d; Imajima, 1966 : 397, fig. 2. a-h; Day, 1967:
274, fig. 12. 10. p-u.

HABITAT. Amongst Phyllochaetopterus socialis tubes at LWM.
RECORD. Matiu Is. - i.

DISTRIBUTION. North Atlantic; South Africa; Mediterranean; Japan; northern
North Pacific.

Exogone uniformis Hartman, 1961

Exogone uniformis Hartman, 1961 : 73, pi. 6, fig. i, pi. 7, figs 1-4; Imajima, 1966 : 400, fig. 4.
a-j.

HABITAT. Coarse sand and silty fine coral sand at LWM.
RECORDS. Komimbo Bay - i ; Matiu Is. - i.
DISTRIBUTION. Southern California; Japan.

Exogone verugera (Claparede, 1868)

Exogone verugera: Fauvel, 1923 : 307, fig. 117. m-r; Imajima, 1966 : 399, fig. 3. a-h; Day, 1967 :
272, fig. 12. 10. g-1.

HABITAT. On Eunice tubifex tube at LWM; on sponge Halichondria sp. at 5 m
depth.

RECORDS. Tetel Is. - i ; Yandina - i.

DISTRIBUTION. North Atlantic; Mediterranean; South Africa; S.W. and E.
Australia; North Pacific.

Sphaerosyllis hirsuta Ehlers, 1897

Sphaerosyllis hirsuta Ehlers, 1897 : 48, pi. 3, figs 58-60; Imajima & Hartman, 1964 : 117, pi. 27,
figs f-1; Imajima, 1966 : 404.

HABITAT. Coarse sand at LWM ; on Eunice tubifex tube at LWM.
RECORDS. Komimbo Bay - i ; Tetel Is. - i.

DISTRIBUTION. Northern North Pacific; Japan; Australia; New Zealand;
southern South America.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 141

Subfamily SYLLINAE
Haplosyllis spongicola (Grube, 1855)

Syllis (Haplosyllis) spongicola: Fauvel, 1923 : 257, fig. 95. a-d; 1953 : 147, fig. 75. a-d; Day,

1967 : 240, fig. 12. i. e-i.
Haplosyllis spongicola: Imajima, i9&6a : 220, fig. 38. a-h.

HABITAT. Abundant in sponges - Neofolitispa dianchora and Halichondria sp. -
from 5 m depth.

RECORDS. Yandina - num.

DISTRIBUTION. Cosmopolitan in tropical and temperate waters.

Langerhansia cornuta (Rathke, 1843)

Syllis (Ehlersia) cornuta: Fauvel, 1923 : 267, fig. 100 g-i; 1953 : 153, fig. 79. g-i.
Syllis (Langerhansia) cornuta: Day, 1967 : 244, fig. 12.2. s-u.
Langerhansia cornuta: Imajima, i966b : 256, fig. 51. a-o.

HABITAT. Coarse sand, silt ; on sponge Halichondria sp. at 5 m depth.
RECORDS. Tetel Is. - 3; Komimbo Bay - 19; Yandina - i.
DISTRIBUTION. Atlantic; Indo-Pacific.

Langerhansia Irosea (Langerhans, 1879)
Langerhansia rosea: Imajima, I966b : 259, fig. 52. a-m.

HABITAT. On serpulid tubes at MTL; coarse sand at LWM.
RECORDS. Tetel Is. - i ; Komimbo Bay - 2.

Syllis longissima Gravier, 1900
Syllis longissima Gravier, 1900 : 154, pi. 9, fig. 7, text-figs. 17-23; Day, 1967 : 243, fig. 12.2. f-i

HABITAT. Crevices in reef platform ; amongst Phyllochaetopterus socialis tubes at
LWM.

RECORDS. Matiu Is. - 8 ; New Manra - i ; Lingatu - 3.

NOTE. This appears to be the first record of S. longissima in the west Pacific
region.

DISTRIBUTION. Red Sea; Persian Gulf; west coast of South America.

142 P. E. GIBBS

Syllis sp. cf. gracilis Grube, 1840

Syllis gracilis: Fauvel, 1923 : 259, fig. 96. f-i; 1953 : 147, fig. 73. f-i; Imajima, ig66a : 248,
fig. 49. a-k; Day, 1967 : 241, fig. 12.1. m-p.

HABITAT. Amongst Phyllochaetopterus socialis tubes at LWM; on sponge
Halichondria sp. at 5 m depth.

RECORDS. Batuona Is. - 3 ; Yandina - i.

Trypanosyllis coeliaca Claparede, 1868

Trypanosyllis coeliaca: Fauvel, 1923 : 270, fig. 101. f-h.

HABITAT. Amongst Phyllochaetopterus socialis tubes at LWM.
RECORD. Batuona Is. - i.

NOTE. Previous records of T. coeliaca are confined to western Europe and the
Mediterranean Sea.

DISTRIBUTION. North Atlantic; Mediterranean.

Trypanosyllis ( Trypanedenta) sp.

HABITAT. Crevices in beachrock at LWM.
RECORDS. Lauvie Is. - 3.

Typosyllis alternata (Moore, 1908)

Typosyllis alternata: Imajima, I966b : 273, fig. 58. a-1.

HABITAT. Chiefly amongst Phyllochaetopterus socialis tubes ; under boulders and
in crevices in reef platform ; on sponge Neofolitispa dianchora at 5 m depth.

RECORDS. Tetel Is. -2; Matiu Is. -5; Cape Esperance-3; Batuona Is. -6;
Yandina - 3.

DISTRIBUTION. Japan; East Indies; west coast of North America.

Typosyllis armillaris (Miiller, 1776)

Syllis (Typosyllis) armillaris: Fauvel, 1923 : 264, fig. 99. a-f; Day, 1967 : 249, fig. 12.4. a-d.
HABITAT. Amongst Phyllochaetopterus socialis tubes at LWM.
RECORDS. Batuona Is. - 2.
DISTRIBUTION. Cosmopolitan.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 143

Typosyllis brachycola (Ehlers, 1897)

Syllis brachycola Ehlers, 1897 : 38, pi. 2, figs 46-47.
Syllis (Typosyllis) brachychola: Augener, 1924 : 362.

HABITAT. Below boulders in silty sand ; amongst Phyllochaetopterus socialis tubes ;
interstices of coral Galaxea.

RECORDS. Tetel Is. - i ; Naro Bay - 2; Batuona Is. - 3; Graham Pt. - 2.

NOTE. T. brachycola is widely distributed in the southern temperate and sub-
antarctic regions but this appears to be the first record from tropical waters.

DISTRIBUTION. Southern South America; New Zealand.

Typosyllis exilis (Gravier, 1900)

Syllis (Typosyllis) exilis Gravier, 1900 : 160, pi. 9, fig. 9, text-figs 28-30; Day, 1967 : 250, fig.
12.4. h-j.

HABITAT. Crevice in reef platform at MTL.

RECORD. Mamara Pt. - i.

DISTRIBUTION. Indo-west-Pacific to Gambier Islands.

Typosyllis lucida (Chamberlin, 1919)

Pionosyllis lucida Chamberlin, 1919 : 8.
Typosyllis lucida: Hartman, 1968 : 489.

HABITAT. Crevices in reef platform.
RECORDS. Kokomtambu Is. - i ; Matiu Is. - 2.

NOTE. This species has not been recorded since it was first described by Chamber-
lin from Laguna Beach, California.

DISTRIBUTION. Southern California.

Typosyllis prolifera (Krohn, 1852)

Syllis (Typosyllis) prolifera: Fauvel, 1923 : 261, fig. 97. a-g; 1953 : 149, fig. 74. a-g; Day, 1967 :

248, fig. 12.3. g-i.
Typosyllis prolifera: Imajima, I9&6b : 292, fig. 65. a-n.

HABITAT. Amongst Phyllochaetopterus socialis tubes at LWM ; on sponge Hali-
chondria sp. at 5 m depth.

RECORDS. Batuona Is. - i ; Yandina - 2.

DISTRIBUTION. North Atlantic; Mediterranean; South Africa; Indo-west-Pacific.

144 P. E. GIBBS

Typosyllis sp.

HABITAT. On sponge Halichondria sp. at 5 m depth.
RECORD. Yandina - i.

Family SPHAERODORIDAE
Sphaerodoridium claparedii (Greeff, 1866)

Sphaerodorum claparedii: Fauvel, 1923 : 379, fig. 149. d-e.
Sphaerodoridium claparedii: Liitzen, 1961 : 415.

HABITAT. On sponge Halichondria at 5 m depth.
RECORD. Yandina - i.

NOTES. This specimen was overlooked during the primary sorting of a collection
of polychaetes from sponges and was included in a large sample of syllids sent to
Dr. M. Imajima who kindly identified it. This record considerably extends the
known distribution of 5. claparedii which hitherto was recorded only from western
Europe.

DISTRIBUTION. European waters.

Family NEREID AE

Subfamily NAMANEREINAE

Cryptonereis n. gen.

Small nereids that are generally similar to Namanereis but which lack frontal
antennae. Prostomium with two pairs of eyes and two biarticulate palps. Proboscis
without paragnaths but with a pair of toothed jaws. Peristomium with three pairs
of tentacular cirri but not parapodia. Parapodia are sesquiramous throughout, each
lacking a notopodial lobe but with a notopodial aciculum. Superior neuropodial
setae are spinigerous, the rest being falcigerous. At maturity, parapodia become
biramous with the development of capillary setae.

TYPE SPECIES. Cryptonereis malaitae Gibbs.

Cryptonereis malaitae n. gen. n. sp.
Fig. 6. (A-F)

DESCRIPTION. Larger specimens measure between 10 and 15 mm long for 55 to
65 segments and are about 2 mm wide.

The prostomium is rounded, roughly semi-circular in shape, with two pairs of eyes

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS

145

and two biarticulate palps with broad palpophores. Frontal antennae are lacking
(fig. 6. A). The peristomium carries three pairs of short tentacular cirri ; on each side
two cirri are attached to a short base which is ventro-lateral in position and the thiid
cirrus arises dorso-laterally on the peristomium (fig. 6. B). The proboscis lacks
paragnaths but carries a pair of brown jaws, each with 9 or 10 teeth and a thin internal
guard (fig. 6. c).

The parapodia are sesquiramous in immature individuals, becoming biramous
over most of the body when mature (see below). Notopodial lobes are lacking but
black notopodial as well as neuropodial acicula persist (fig. 6. D). Neuropodia are
bilobed with pointed pre-setal and rounded post-setal lips. The dorsal cirri are
short and stout and the ventral cirri are very small.

The neuropodial setae consist of one or two spinigers with minutely serrated
blades (fig. 6. E) and four or five falcigers with short ciliated blades (fig. 6. F). The
shaft and blade of the superior falciger often appear to be fused but the articulations
of the other falcigers are usually distinct.

Most of the specimens appear to be approaching maturity and, on preservation,
gametes have been released from the parapodia at points just above the dorsal cirri.

FIG. 6. Cryptonereis malaitae n.gen., n.sp. (A-B) Dorsal and lateral views of anterior
region, (c) Left jaw in ventral view. (D) Parapodium from middle body of mature
specimen. (E) Spiniger. (F) Falciger.

146 P. E. GIBBS

In these specimens, long, slender capillary setae are present in both rami of the
parapodia from setigers 8 or 9 to the last few segments. In some, these capillary
setae are short and few in number but in others, probably at a more advanced stage
of maturity, they are very much longer, being about three times the length of the
falcigers. In the latter specimens, each parapodium has a compact notopodial
bundle, composed of about 20 capillaries, arising from between the dorsal cirrus
and the notopodial aciculum, and a further 8 to 10 capillaries appear amongst the
neurosetae (fig. 6. D). The specimen chosen for the holotype shows this condition.

In the collection there is an immature individual which is only 6-5 mm long for
35 segments (incomplete). This specimen does not possess the capillary setae found
in the larger specimens. It is thought therefore that C, malaitae has an epitokous
phase at maturity in which capillary (or natatory) setae (recalling those that are
formed in the epitokes of some Syllidae) are developed and that the majority of the
type specimens, collected on 20.xi.65, are approaching, or at, this stage of develop-
ment.

C. malaitae is apparently dioecious. The oocytes are yolky and those released on
preservation by the females vary greatly in size but are mainly between 100 to 150(1
in diameter. Spermatozoa are recognizable in the coelomic contents of the more
mature males. In life, specimens are purplish but assume a brownish colour in
alcohol.

It seems inadvisable to enlarge the generic description of the relatively well-known
and widespread genus Namanereis Chamberlin (see Hartman, 19590) to include
those forms which lack frontal antennae, hence the need to erect a new genus. Apart
from its lacking frontal antennae and also its possession of capillary setae at maturity,
Cryptonereis closely resembles Namanereis and the relationship between these two
genera parallels that between Micronereis Claparede and Micronereides Day in terms
of the presence or absence of antennae.

HABITAT. Between the fibres composing a leaf frond of the coconut palm found
stranded at about MTL.

RECORDS. Alite Harbour, Langa-Langa Lagoon (Malaita) - 14.
British Museum (Natural History) Registration No. Holotype 1970-30

Paratypes 1970-31

NOTES. It is difficult to decide whether the habitat of C. malaitae is marine or,
like many other Namanereinae, damp-terrestrial since either is possible, depending
on the period of time the leaf frond had been immersed in seawater. Interestingly,
further specimens of this, or a closely related species, were collected by the Expedition
Land Party (Mr. J. Peake and Dr. J. Greenslade) in moist leaf litter at an altitude
of about 350 m at two separate localities, namely Betimatu on Mt. Austen, Guadal-
canal and on Kolombangara in the New Georgia Group. This material consists of
two specimens (one incomplete and the other dried) both about 5 mm long and
resembling immature C. malaitae in detail. However in both specimens the pros-
tomium is damaged and because the presence or absence of frontal antennae cannot
be determined their identity must remain in question. It is possible that these two

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 147

specimens could be Namanereis amboinensis (Pflugfelder, 1933, as Lycastopsis
amboinensis) which resembles the immature stage of C. malaitae in all details but
possesses antennae.

Namalycastis indica (Southern, 1921)

Lycastis indica Southern, 1921 : 578, pi. 19, fig. 2. a-j, text-fig. 2. a-d.
Namalycastis indica: De Silva, I965a : 5, fig. 2. a-e; Day, 1967 : 301, 14.2. p-s.

HABITAT. Burrowing within the fibrous husks of Nipa palm nuts immersed in
brackish water (5'6% ).

RECORD. Lagoon at the mouth of the Lunga R. at Lunga Pt. - 12.

NOTES. N. indica was found only in those Nipa nuts which were waterlogged,
lying along and just below the water-line of the swamp. The largest specimen is
60 mm long.

DISTRIBUTION. Indian Ocean.

Subfamily NEREINAE
Ceratonereis costae (Grube, 1840)

Ceratonereis costae : Fauvel, 1923 : 349, fig. 136. a-f; 1953, X 94 % 98. a-f; Day, 1967 : 325, fig.
14.10. h-1.

HABITAT. Amongst Phyllochaetopterus socialis tubes; on Halichondria sp. at
5 m depth.

RECORDS. Batuona Is. - i ; Yandina - i.

DISTRIBUTION. North Atlantic; Mediterranean; Indo-west-Pacific.

Ceratonereis erythraeensis Fauvel, 1918

Ceratonereis erythraeensis Fauvel, 1918 : 505, fig. 2. a-k; Day, 1967 : 327, fig. 14.10. o-t.

HABITAT. Muddy silt under boulders at MTL.

RECORDS. Naro Bay - 20.

DISTRIBUTION. Indian Ocean; East Australia; Japan.

148 P. E. GIBBS

Ceratonereis mirabilis Kinberg, 1866

Ceratonereis mirabilis: Gravier, 1901 : 172, pi. n, fig. 42; Fauvel, 1953 : 200, fig. 103. a-c; Day,
1967 : 324, fig. 14.10. a-g.

HABITAT. Silty sand under boulders; in Acropora rubble; within interstices of
Galaxea; on Halichondria sp. at 5 m depth.

RECORDS. Tetel Is. - 2 ; Komimbo Bay - i ; Graham Pt. - 2 ; Yandina - 3.
DISTRIBUTION. Circumtropical.

Nereis (Neanthes) caudata (Delle Chiaje, 1825)

Nereis cricognatha: Fauvel, 1953 : 180, fig. 91. a-c.

Nereis (Neanthes) arenaceodonta: Pettibone, 1963 : 162, figs 44.!, 45.6.

Nereis (Neanthes) caudata: Day, 1967 : 321, fig. 14.9. f-j.

HABITAT. Under coral boulder in muddy silt; sand at 5 m depth.
RECORDS. Komimbo Bay - i ; ML 229 - i.
DISTRIBUTION. Cosmopolitan.

Nereis (Neanthes) unifasciata Willey, 1905

Nereis (Neanthes) unifasciata Willey, 1905 : 271, pi. 4, figs 85-88; Fauvel, 1953 : 182, 92. a-h;
Day, 1967 : 318, fig. 14.7. u-y.

HABITAT. Under coral boulder; in beachrock.
RECORDS. Komimbo Bay - i ; Lauvie Is. - i.
DISTRIBUTION. Tropical Indo-west-Pacific.

Nereis (Neanthes) sp. cf. kerguelensis Mclntosh, 1885

Nereis kerguelensis Mclntosh, 1885 : 225, pi. 35, figs 10-12, pi. i6A, figs 17-18.

HABITAT. Silty sand in crevice of Porites boulder.
RECORD. Graham Pt. - i.

NOTES. The specimen corresponds to N. kerguelensis in all details except that the
notopodia of the anterior segments lack intermediate pre-setal lobes, giving a bi-
lobed, not tri-lobed structure.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS

149

Perinereis cultrifera (Grube, 1840)
(Fig. 7. A)

Perinereis cultrifera : Fauvel, 1923 : 352, fig. 137. a-1; 1953 : 206, fig. 106. a-1; Day, 1967 : 337,
fig. 14.13. o-q.

HABITAT. Chiefly under coral boulders on sand at MTL; also in crevices in reef
platform and in beachrock.

RECORDS. Kokomtambu Is. -i; Tetel Is. -3; Matiu Is. -6; New Manra-32;
Maraunibina Is. -4; Lauvie Is. -i; Graham Pt. -4; Yandina-i; Fanalei-2
(heteronereids).

NOTES. Two female heteronereid stages of this species are included in the
collection made during the 'palolo' rising at Fanalei on 2.xi.66 (local name - 'falisi
ogu'). They were taken between 2000 and 2200 hours and correspond to the
description of this stage given by Izuka (1912) and Fauvel (1923) except that lower
ligule of the notopodium in the modified parapodium has a small ear-shaped lobe
arising from its ventral margin (fig. 7. A). These specimens measure 45 and 52 mm
long for 116 and 225 segments respectively, with the modified parapodia commencing
at setigers 20 to 22.

DISTRIBUTION. Cosmopolitan.

1.0mm

FIG. 7. Perinereis cultrifera (A) Modified parapodium of heteronereid (?) stage (setae
omitted). Perinereis nuntia (B) Modified parapodium of heteronereid (<$) stage (setae
omitted) .

150 P. E. GIBBS

Perinereis nigropunctata (Horst, 1889)

Nereis (Perinereis) nigro-punctata Horst 1889 : 171, pi. 8, figs 1-3; 1924 : 171.

Perinereis nigro-punctata: Fauvel, 1953 : 210, fig. 107. b-f; Day, 1967 : 337, fig. 14.13. r-v.

HABITAT. Amongst Phyllochaetoptems socialis tubes; under coral boulders;
abundant in beachrock.

RECORDS. Batuona Is. - 2 ; Graham Pt. - 2 ; Lauvie Is. - num.
DISTRIBUTION. Tropical Indo-west-Pacific.

Perinereis nuntia (Savigny, 1818)

(Fig. 7. B)
Perinereis nuntia: Fauvel, 1953 : 212, fig. 109. a-g; Day, 1967 : 334, fig. 14.12. p-s.

HABITAT. Chiefly found under boulders lying on clean sand and gravel at MTL.

RECORDS. Haroro - 3 ; Tetel Is. - 3 ; Cape Esperance - 17 ; Matiu Is. - 2 ; New
Manra - 3 ; Mouth of Lunga R. - 4 ; Yandina - 50 ; Honiara - 20 ; Vera Vera Entrance
- i (heronereid ($} ).

NOTES. In these specimens there are I to 3 paragnaths in group i and the parag-
naths of group VI are usually flattened, sometimes mixed with conical forms.
However the number of paragnaths in group V is very variable : for example, in the
sample of 50 individuals from the Yandina population, one specimen has four
paragnaths in group V, 26 have three arranged in a triangle, n have two and 12
have only one. Thus in terms of the subspecific forms (see Fauvel, 1953) Perinereis
nuntia brevicirrus composes about half the population and P. nuntia vallata about a
quarter, the remaining quarter being intermediate in character between these two
forms.

The male heteronereid stage of P. nuntia brevicirrus was captured swimming near
the surface at Vera Vera Entrance on 8.xi.65 at 1500 hours. It measures 90 mm
long for about 150 segments, the modification of the parapodia starting between
setigers 23 to 27. The structural details of the modified parapodium (fig. 7. B) follow
those described and figured by Izuka (1912, as N. mictodonta).

DISTRIBUTION. South Africa; Indo-Pacific.

Platynereis insolita Gravier, 1901

Platynereis insolita Gravier 1901 : 197, pi. 12, fig. 53, text-figs 203-206; Day, 1967 : 307, fig.
14.4.1.

HABITAT. Under coral boulder; amongst Phyllochaetopterus socialis tubes and
Amphiroa growth.

RECORDS. Matiu Is. - i ; Komimbo Bay - i ; New Manra - 2.
DISTRIBUTION. Indian Ocean.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 151

Pseudonereis anomala Gravier, 1901

Pseudonereis anomala Gravier, 1901 : 191, pi. 12, figs 50-52, text-figs 194-202; Fauvel, 1953 :
217, fig. no. e-g; Day, 1967 : 333, fig. 14.12. g-j.

HABITAT. Abundant amongst Phyllochaetopterus socialis tubes and Amphiroa;
burrowing in reef platform and beachrock.

RECORDS. Mamara Pt. - i ; Batuona Is. - num. ; New Manra - num. ; Yandina -
6 ; Maramasike Pg. - 3.

DISTRIBUTION. Indo-Pacinc.

Pseudonereis masalacensis (Grube, 1878)

Nereis (Lycoris) masalacensis Grube, 1878 : 75, pi. 5, fig. 4.
Pseudonereis masalacensis: Fauvel, 1947 : 51, fig. 49. a-k.

HABITAT. Under coral boulders on sand at MTL; amongst Phyllochaetopterus
socialis tubes and algal growth (Amphiroa).

RECORDS. Batuona Is. - 2 ; New Manra - 3 ; Yandina - 2.
DISTRIBUTION. West and South Pacific.

Pseudonereis variegata (Grube, 1857)

Pseudonereis gallapagensis : Fauvel, 1953 : 2I 5. fig- IIQ - a ~ c -
Pseudonereis variegata: Day, 1967 : 331, fig. 14.12. a-f.

HABITAT. Burrowing in reef platform and beachrock; amongst Amphiroa; under
boulders on clean sand.

RECORDS. Matiu Is. - 8 ; Sifola - 8 ; New Manra - 2 ; Lauvie Is. - 3 ; Yandina - 2.
DISTRIBUTION. Circumtropical.

Solomononereis n. gen.

Prostomium with two antennae, two biarticulate palps and two pairs of eyes.
Peristomium with four pairs of tentacular cirri, dorsal pairs long, ventral pairs much
shorter. Proboscis with small, rod-like chitinous paragnaths arranged in eight
discrete groups on the maxillary ring; oral ring without paragnaths. Parapodia
biramous with spinigerous setae in both rami throughout the body; falcigerous setae
in both rami in the middle and posterior segments.

TYPE SPECIES. Solomononereis marauensis Gibbs

152

P. E. GIBBS

Solomononereis marauensis n. gen. n. sp.

(Fig. 8. A-H)

DESCRIPTION. The holotype measures 45 mm for 123 segments and is incomplete
posteriorly. The largest of the three paratypes is 40 mm for 157 segments. The
width of the body is about 3 mm.

The prostomium is rectangular in shape with two short, widely spaced antennae,
two pairs of eyes, and a pair of palps with stout palpophores, often with a constriction
just below the tip, and small palpostyles (fig. 8. A). The peristomium is weakly
developed and is present as a narrow band on the dorsal surface but does not form a
distinct ring ventrally. Of the four pairs of tentacular cirri, the two dorsal pairs
are very long, extending back as far as setiger 20 but the ventral pairs are short,
reaching to setiger 3 only.

FIG. 8. Solomononereis marauensis n.gen., n.sp. (A) Dorsal view of anterior region and
proboscis. (B) Ventral view of proboscis, (c) Paragnaths of Group II. (D-F) Anterior,
middle and posterior parapodia. (G) Notopodial falciger. (H) Subacicular neuro-
podial spiniger.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 153

The proboscis carries a pair of brown, curved jaws, each with 7 or 8 teeth.
The chitinous paragnaths are confined to the maxillary ring ; they are minute, rod-like
(7-iO{/. in diameter) structures which are closely packed together in eight groups,
three on the dorsal side and five on the ventral (fig. 8. A, B). Each of these discrete
groups is composed of 10 to 15 paragnaths (fig. 8. c).

The structure of the parapodia changes progressively throughout the body and
those of the anterior region are larger than those of the middle and posterior parts of
the body. In the anterior region, the notopodium has two pointed ligules and the
neuropodium has a bilobed upper ligule, composed of a pointed pre-setal lip and a
rounded post-setal lip, and a pointed lower ligule (fig. 8. D). Between setigers 20 and
30, the upper notopodial ligule and the lower neuropodial ligule decrease in size,
the former being lost altogether in more posterior segments while the latter persists
as a rudiment (fig. 8. E, F). The dorsal cirri are slightly longer than the parapodial
lobes in anterior segments but are relatively much longer in the posterior segments
because of the diminished size of the parapodia.

At the base of the dorsal cirri, there are conspicuous glands which are roughly
oval in shape and have a yellowish colour in preserved (alcohol) specimens. Each
gland consists of fascicles of spindle-shaped cells which appear to open to the exterior
at a pore situated just above the base of the dorsal cirrus.

Spinigerous setae are present in both notopodial and neuropodial bundles through-
out the body. The notosetae are all homogomphic while the neurosetae vary from
homogomphic to hemigomphic in the superior position, becoming heterogomphic
in the inferior part. Superior sub-acicular spinigers have coarsely serrated blades
(fig. 8. H) but all others are finely serrated. In the region of setiger 40, and persisting
to the posterior end, two or three homogomph falcigers (fig. 8. G) appear in the
notopodial bundle and, in addition, a single heterogomph falciger appears in the
supra-acicular part of the neuropodium. Each falciger has an elongate, ciliated
blade with a curved tip supported by a distinct tendon. All setae have a crystalline
appearance.

A dark brown pigment remains over the dorsal surface of the proboscis, prostomium
and anterior segments in preserved specimens.

Solomononereis resembles Ceratonereis in having paragnaths only on the maxillary
ring of the proboscis but this similarity is superficial, the shape, as well as the arrange-
ment, of the 'toothpick' or rod-shaped paragnaths possessed by Solomononereis
being very different to that of the conical paragnaths found in Ceratonereis. Other
features separating Solomononereis from other nereid genera include the weakly
developed peristomial ring and the arrangement of the setae, i.e. spinigers are
present in both rami throughout the body and, in addition, a few falcigers appear in
both rami of the middle and posterior parapodia.

HABITAT. In sticky mud above MTL close to freshwater outflows.

RECORDS. Fintry Pt. - i (holotype) ; Nggela Is. (Sandfly Passage) - 3.
British Museum (Natural History) Registration No. Holotype 1970-32

Paratypes 1970-33

154 p - E - GIBBS

Family NEPHTYIDAE
Nephtys (Aglaophamus) munamaorii n. sp.

(Fig. 9. A-B)

DESCRIPTION : The holotype (from ML 37) is the largest specimen in the col-
lection. It measures 16 mm in length for 60 segments and is about i mm wide.

The prostomium is rectangular in shape, its length being about i -5 times the width,
and has four small antennae on the antero-lateral margins. The proboscis has a
long median papilla on its dorsal side with 14 rather indistinctly aligned rows of
5 or 6 papillae. Two small eyes, which appear as black spots, are situated near the
posterior border of the second segment.

Branchiae commence on setiger 3 and are well developed in the anterior half of the
body but gradually diminish in size in the posterior half so as to be absent from the
last 10 or 12 segments. Each branchia is involute with a dorsal cirrus at its base
(fig. 9. A).

The setigerous lobes of the parapodial rami are pointed. In the anterior segments,
both the pre-setal and the post-setal lamellae are rounded, the latter being slightly
larger, but in posterior segments these lamellae are much reduced in size (fig. 9. B).
There are no superior neuropodial lamellae.

Setae are of the three usual types. Barred (or laddered) -capillary setae compose
the anterior fan which also includes a few smooth capillaries : in posterior segments
the barred setae are longer and more slender with less distinct bars. Smooth or
minutely serrated capillaries form the posterior fan : in the latter, there is also a row
of lyriform forked setae.

Four species belonging to the subgenus Aglaophamus have been described from the
tropical Indo-west-Pacific region. These are N. (A.) dibranchis Grube, N. (A.)

0.5mm

FIG. 9. Nephtys (Aglaophamus) munamaorii n.sp. (A-B) Anterior views of parapodia
from setigers 19 and 45 respectively (setae omitted).

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 155

lyrochaeta Fauvel, N. (A.) mirasetis Hoagland, and N. (A.) sinensis Fauvel. In all
four species the neuropodia of anterior segments have superior lamellae. The
absence of this lamella, in addition to further differences in the parapodial structure,
particularly the shape of the pre- and post-setal lamellae, is sufficient to distinguish
N. (A.) munamaorii.

HABITAT. Mud and silty sand, 2-22 m depth.

RECORDS. ML 37-2 (including holotype) ; ML 69 - i ; ML 157 - i ; ML 188 - 8;
ML 194 - 2 ; ML 196 - i ; ML 218 - 2.
British Museum (Natural History) Registration No. Holotype 1970-34

Paratypes 1970-35-41

Nephtys (Micronephthys) sphaerocirrata Wesenberg-Lund, 1949

Nephthys sphaerocirrata Wesenberg-Lund, 1949 : 294, figs 24-26.
Nephtys (Micronephthys) sphaerocirrata: Day, 1967 : 347, fig. 15.3. a-d.

HABITAT. Silty sand at 15 m depth.

RECORD. ML 116 - i.

NOTES. The single specimen is an anterior fragment of 33 segments, 5 mm in
length. It differs from the type specimen in having two pairs of reddish eyes situated
near the posterior margin of the prostomium. These may represent a sexual
character since the specimen is a mature female containing oocytes. Other details
agree with the type description.

DISTRIBUTION. Persian Gulf; South Africa; Marshall Islands.

Nephtys (Nephtys) sp. cf. palatii Gravier, 1904
Nephthys palatii Gravier, 1904 : 472; 1906 : 129, pi. i, figs 163-164, text-figs 286-289.

HABITAT. A wide variety of deposits from mud to coarse sand, 2-18 m depth.

RECORDS. ML 39 - i ; ML 68 - i ; ML 97 - i ; ML 156- i ; ML 191 - i ; ML 192 - 1 ;
ML 196 - i ; ML 203 - i ; ML 229 - 2.

NOTE. These specimens are close to N. palatii but the identity is uncertain
because, on dissection, a median dorsal papilla could not be seen on the proboscis.

Family LACYDONIIDAE
Paralacydonia weberi Horst, 1923

Paralacydonia weberi Horst, 1923 : 221, figs 1-2; Fauvel, 1953 : 129, figs 65. e-f.
HABITAT. Mud and sand, 2-24 m depth.

156 P. E. GIBBS

RECORDS. ML 29 - i ; ML 190 - i ; ML 191 - i ; ML 230 - i.

NOTE. The largest of the specimens is 10 mm long for 50 segments and is much
smaller than the holotype (40 mm for 100 segments) which was dredged from deep
water (959 m).

DISTRIBUTION. Burma; East Indies; Samoa; Bass Strait.

Family GLYCERIDAE

Subfamily GLYCERINAE

Glycera gigantea Quatrefages, 1865

Glycera gigantea : Fauvel, 1923 : 387, fig. 152. d-k; 1953 : 296, fig. 152. d-k; Day, 1967 : 362,
fig. 16.2. 1-n.

HABITAT. A wide variety of deposits, chiefly silty mud and sand, from above
MTL to 33 m depth.

RECORDS. Tetel Is. -i; Florida Is. -3; Komimbo Bay -7; New Manra-i;
Graham Pt. -8; Fintry Pt. -4; Maraunibina Is. -i; ML 229-1. (ML stations
100, no, 192, 195 - one juvenile (?) at each).

DISTRIBUTION. North Atlantic; Mediterranean; Laccadive Sea; New Britain;
N.E. Australia; Japan; eastern North Pacific.

Glycera lancadivae Schmarda, 1861

Glycera lancadivae Schmarda, 1861 : 95, with text-figs; Fauvel, 1953 : 291, fig. 147. g-h; Day,
1967 : 359-

HABITAT. A wide variety of deposits, from fine coral mud to coarse sand and
shell gravel from above MTL to n m depth.

RECORDS. Tetel Is. -3; Komimbo Bay -4; Naro Bay-i; Matiu Is. -11;
Maraunibina Is. - 22 ; Graham Pt. - 8 ; Fintry Pt. - 9 ; Yandina - 4; ML 98 - i ;
ML 203 - i ; ML 229 - 2.

DISTRIBUTION. Tropical Indian Ocean; N.E. Australia; New Caledonia.

Glycera longipinnis Grube, 1878

Glycera longipinnis Grube, 1878 : 182, pi. 8, fig. 9; Fauvel, 1932 : 125, pi. 4, figs 11-14; I 953 :
291, fig. 148. a-d; Day, 1967 : 356, fig. 16.1. a-f.

HABITAT. Muddy silt (Thalassia flat) at LWM.
RECORD. Komimbo Bay - i.
DISTRIBUTION. Tropical Indo-west-Pacific.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 157

Glycera rouxi Audouin & Milne-Edwards, 1833

Glycera rouxi: Fauvel, 1923 : 389, fig. 153. a-c; 1953 : 2 97. fig- J 49- a -d; Day, 1967 : 362, fig.
16.3. a-d.

HABITAT. Silty mud at 9 m depth.
RECORD. ML 38-1.

DISTRIBUTION. North Atlantic; Mediterranean; Indo-west-Pacific to Japan;
California.

Family EUNICIDAE
Subfamily EUNIGINAE
Eunice afra Peters, 1854

Eunice afra: Grassland, 1904 : 289, pi. 20, fig. 1-5, text-figs 43-45; Fauvel, 1953 : 2 35> fig- IJ 6.
h-i; Day, 1967 : 392, fig. 17.5. a-e.

HABITAT. Boring in coral on reef platform and in beachrock; amongst Phyl-
lochaetopterus socialis tubes at LWM.

RECORDS. Matiu Is. -4; Mamara Pt. -2; Batuona Is. -2; New Manra-i;
Lauvie Is. - 2; Lingatu - 4; Yandina - i ; Maramasike Pg. - 5.

DISTRIBUTION. Indo-west-Pacific.

Eunice antennata (Savigny, 1820)

Eunice antennata: Crossland, 1904 : 312, pi. 22, figs 1-7, text-figs 56-60; Fauvel,! 953 : 240,
fig. 118. f-g; Day, 1967 : 384, fig. 17.2. k-q.

HABITAT. Boring in coral and under coral boulders on reef platform.
RECORDS. Matiu Is. - 5 ; Graham Pt. - 2 ; Yandina - i.
DISTRIBUTION. North Atlantic; tropical Indo-Pacific.

Eunice aphroditois (Pallas, 1788)
Eunice aphroditois: Fauvel, 1953 : 233, fig. 117. a-g; Day, 1967 : 389, fig. 17.4. l-o.

HABITAT. In Acropora rubble and under coral boulder on reef platform at LWM.
RECORDS. Tetel Is. - i ; Maraunibina Is. - 2.

DISTRIBUTION. North Atlantic; Mediterranean; Indo-Pacific to Japan and
southern California.

158 P. E. GIBBS

Eunice coccinea Grube, 1878

Eunice coccinea Grube, 1878 : 153, pi. 9, fig. i; Grassland, 1904 : 297, pi. 20, figs 6-7, text-figs
46-51; Fauvel, 1953 : 2 36, fig- 118. a-e; Day, 1967 : 389.

HABITAT. Boring in coral on reef platform.

RECORDS. Komimbo Bay - i ; Mamara Pt. - i ; New Manra - i ; Lingatu - i ;
Maramasike Pg. - 2; Fanalei - 2 (posterior fragments).

NOTES. The two posterior fragments in the collection from the Fanalei 'palolo'
rising (2-3.ix.66) can be identified by their relative size, absence of gills, parapodial
structure and setae. One is female and contains large spherical oocytes (360(0. in
diameter). On one side of each of these oocytes there is a patch of dark green
pigment, surrounding a clear circular spot. The time of spawning is recorded as
midnight to 0600 hours. The local name is 'raka-raka'.

DISTRIBUTION. Gulf of Guinea; tropical Indo-west-Pacific.

Eunice grubei Gravier, 1900

Eunice grubei Gravier, 1900 : 258, pi. 14, figs. 87-88, text-figs 125-129; Fauvel, 1953 : 237, fig.
119. a-e; Day, 1967 : 391.

HABITAT. Sedimented crevices in Forties boulders.
RECORDS. Graham Pt. - 2.

NOTES. One specimen is complete and, in this, the branchiae, which are present
only on the anterior half of the body, have a maximum of 4 or 5 filaments.

DISTRIBUTION. Tropical Indo-west-Pacific.

Eunice marovoi n. sp.

(Fig. 10. A-H)
Eunice sp.: Gibbs, 1969 : 455, fig. 133.

DESCRIPTION. Complete specimens from the dredged material have a maximum
length of 15 mm for 60 to 70 segments and a width of i-o to 1-5 mm. Larger
specimens discovered living on Cerithium vertagus are all incomplete but are
estimated to have been 20 to 25 mm in length. The holotype from ML 96 is 15 mm
long for 66 segments.

The prostomium is bilobed with a shallow median groove between the palps
(fig. 10. A). The antennae are weakly annulated and the median is about twice
as long as the laterals, which, in turn, are about equal in length to the tentacular cirri
(fig. 10. B). The eyes are black and reniform in shape.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS

159

The mandibles are elongate calcified plates. The maxillae are almost transparent
but the toothed edges are brownish in colour and the medial and anterior edges of
the maxillae supports ("carriers") are black, forming a T-shaped figure (fig. 10. c).
Due to the transparent nature ot the maxillae the details are difficult to determine.
Maxillae I are falcate and devoid of basal teeth. The left Maxilla II is falcate with
4 to 6 ill-defined basal teeth. The right Maxilla II, left Maxilla III and Maxillae IV
all have 10 to 13 small teeth while Maxillae V are toothless plates. The maxillary
formula thus reads: MX. 1 = 1 + 1; MX. II = (i + (4 6)) + (10 13) ; MX. Ill
= (10 13) + o; MX. IV = (10 13) + (10 13).

Branchiae commence on setiger 3 on all specimens and these are strongly arched
over the dorsum, with a maximum of 5 or 6 filaments (fig. 10. D). In the larger
specimens there are 21 to 23 pairs (i.e. extending to setigers 23 to 25) but smaller

FIG. 10. Eunice marovoi n.sp. (A) Dorsal view of prostomium. (B) Antero-lateral view
of anterior region, (c) Jaws. (D-E) Branchia and parapodium of setiger 10. (F) Comb
seta. (G) Falciger. (H) Acicular seta.

160 P. E. GIBBS

specimens have only 15 to 20 pairs. The dorsal cirri are shorter but slightly stouter
than the branchial filaments. On branchiferous segments, the ventral cirri are
short and are swollen at their bases (fig. 10. E), a feature that is absent in the more
posterior segments.

Acicula and setae are transparent. The capillary setae are slender with finely
serrated edges. Comb setae are few in number and have about 7 teeth, one of the
outer pair being greatly elongated (fig. 10. F). The falcigerous setae have bidentate
blades with faintly serrated guards (fig. 10. G). Acicular setae are tridentate with
rounded guards (fig. 10. H) ; two or three occur in each parapodium from setigers 15
to 20 to the posterior end. The projecting tips of the acicula are rounded and slightly
curved.

The colour of preserved specimens is white. Despite their small size, a number of
the specimens are mature with coelomic gametes. It would appear therefore that
E. marovoi does not exceed 20 to 25 mm in length even when fully grown.

E. marovoi is distinct from all other Eunice species known from the Indo-west-
Pacific region in terms of its maxillary structure, the number of branchiae and
branchial filaments, in combination with the structure of the setae.

HABITAT. A wide variety of deposits ranging from mud to coarse sand from
LWM to 33 m depth. At LWM on a sand flat to the north of Tankomo Is. (in the
southern part of Marovo Lagoon) E. marovoi was discovered attached to the shell
of the gastropod Cerithium vertagus L.

RECORDS. ML 37 - i ; ML 96-1 (holotype) ; ML 97 - i ; ML 100 - i ; ML no - 9 ;
ML 134 - i ; ML 192 - 2 ; ML 218 - i ; ML 229 - i ; ML 230 - 2 ; north of Tankomo

Is. -5.

British Museum (Natural History) Registration No. Holotype 1970-42

Paratypes 1970-43-52

NOTES. The tube of this species is constructed of coarse bottom material, such
as scaphopod and bivalve she)l fragments, cemented to an organic lining. In those
specimens associated with Cerithium vertagus, the tube was constructed along the
length of the upper surface of the gastropod shell (see Gibbs, 1969).

Eunice norvegica (Linnaeus, 1767)

Eunice floridana : Fauvel, 1923 : 402, fig. 157. a-g; 1953 : 235, fig. 117. a-g.
Eunice norvegica: Pettibone, 1963 : 240, fig. 63. f; Day, 1967 : 388, fig. 17.3. r-v.

HABITAT. On surface of Acropora sp.

RECORD. Yandina - 1.

NOTES. The specimen is 17 mm long, 3 mm wide and has 70 segments (apparently
regenerating from setiger 27). Its preserved colour is medium brown with three
whitish spots per segment arranged in longitudinal lines down the body; the gills

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 161

and dorsal cirri are purple. In life the colour was bluish-black with orange spots.
The maxillary formula is: MX. I = i -|- i; MX. II = 7 -j- 7; MX. Ill = 6 + o;
MX. IV = 4 + 8; MX. V = i + i. The labrum is calcined and pearly- white in
colour. The branchiae start on setiger 5 and have a maximum of 6 or 7 filaments.
Acicular setae are black with conspicuous guards.

In specimens from other localities the branchiae commence between setigers 7 to
10 but otherwise this specimen agrees in detail. Apart from a single specimen
from Indo-China (Fauvel, 1939) no other records of E. norvegica from the Pacific
Ocean have been traced despite the species being widespread and occurring in depths
of over 1500 m (Pettibone, 1963).

DISTRIBUTION. North Atlantic ; Mediterranean ; Indian Ocean ; Indo-China.

Eunice tentaculata Quatrefages, 1865

Eunice tentaculata: Fauvel, 1953: 234, fig. 118. m-p; Day, 1967 : 391, fig. 17.4. s-v.

HABITAT. Under coral boulders on silty sand ; amongst Phyllochaetoptems socialis
tubes.

RECORDS. Maraunibina Is. - 2; Batuona Is. - i.
DISTRIBUTION. Indo-west-Pacific.

Eunice tubifex Grassland, 1904

Eunice tubifex Grassland, 1904 : 303, pi. 21, figs 1-8, text-figs 52-55; Fauvel, 1953 : 2 3 2 n g- Il( >.
a-g; Day, 1967 : 386, fig. 17.3. k-q.

HABITAT. In coral at LWM and on wharf piles at 5 m depth.
RECORDS. Tetel Is. - 2 ; Yandina - 2.
DISTRIBUTION. Tropical Indo-west-Pacific.

Eunice (Palola) siciliensis Grube, 1840

Eunice (Palola} siciliensis: Fauvel, 1923 : 405, fig. 159. e-m; 1953 : 241, fig. 121. e-m; Day, 1967 :
382, fig. 17.2. a-f.

HABITAT. Boring in coral and beachrock; in Acropora rubble and shell gravel;
under coral boulders in silty sand.

RECORDS. Kokomtambu Is. - i ; Tetel Is. - 16 ; Komimbo Bay - 2 ; Mamara Pt. -
3 ; Matiu Is. - 34 ; Yandina - 2 ; Lingatu - 6 ; New Manra - 9 ; Lauvie Is. - n ;
Graham Pt. - 4; Maramasike Pg. - 10; Fanalei - 4 (posterior fragments).

162 P. E. GIBBS

NOTES. The four posterior fragments collected during the spawning at Fanalei
can be readily identified by the absence of comb and acicular setae, the black
acicula and the dark spot on the ventral surface of each segment. Three of the
fragments are female and contain spherical oocytes about 2OO[A in diameter. The
time of spawning on 2.xi.66 was between 2000 and 2200 hours. The local name is
"falisi ogu".

DISTRIBUTION. Mediterranean; circumtropical.

Lysidice collaris Grube, 1870
Lysidice collaris: Fauvel, 1953 : 248, fig. 124. a-g; Day, 1967 : 402, fig. 17.8. a-f.

HABITAT. Boring in coral and beachrock; amongst Phyllochaetopterus socialis
tubes ; in the sponge Halichondria sp. at 5 m depth.

RECORDS. Matiu Is. - 5 ; Batuona Is. - 1 ; New Manra - 3 ; Sifola - 2 ; Lingatu - 2 ;
Mamara Pt. - 2 ; Graham Pt. - i ; Lauvie Is. - i ; Yandina - 12 ; Fanalei - i.

NOTES. The specimen from the Fanalei "palolo" rising is spent. It has enlarged
eyes provided with lenses.

DISTRIBUTION. Tropical Indo-Pacific to Japan and Gambier Islands.

Marphysa macintoshi Crossland, 1903

Marphysa macintoshi Crossland, 1903 : 137, pi. 14, figs 3-6, text-fig. 12; Fauvel, 1953 : 246;
Day, 1967 : 396, fig. 17.6. a-e.

HABITAT. Silty mud at MTL.
RECORDS. Nggela Is. - 3.
DISTRIBUTION. Tropical Indian Ocean.

Nematonereis unicornis (Grube, 1840)

Nematonereis unicornis: Fauvel, 1923 : 412, fig. 162. h-n; 1953, 249, fig. 124. h-n; Day, 1967 : 403,
fig. 17.8. j-n.

HABITAT. Under boulders in muddy silt at MTL; amongst Phyllochaetopterus
socialis tubes at LWM.

RECORDS. Tetel Is. - i ; Batuona Is. - i.

DISTRIBUTION. North Atlantic; Mediterranean; tropical Indo-west-Pacific.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 163

Subfamily ONUPHINAE
Onuphis (Nothria) holobranchiata Marenzeller, 1879

Onuphis (Nothria] holobranchiata Marenzeller, 1879 : 132, pi. 4, fig. i; Izuka, 1912 : 106, pi. n,
figs 10-12; Fauvel, 1953 : 2 56, fig- 127. f-h; Day, 1967 : 424, fig. 17.13. f-g.

HABITAT. A wide variety of deposits, from silty sand to coarse sand and shell
gravel from MTL to LWM ; mud with organic debris at 2 m depth.

RECORDS. Tetel Is. -14; Komimbo Bay-i; Matiu Is. -i; Kalota Is. -i;
ML 68 - 37.

NOTES. The distribution of the simple branchiae over the anterior segments
shows some degree of variation, perhaps reflecting differences in the growth stages.
For example, in the smaller specimens present in the large sample obtained at ML 68
the branchiae commence on setigers 2 to 5 whereas in the larger specimens from the
same sample they begin on the first setiger, as typical for the species. Specimens
of all sizes possess similar pseudocompound hooks.

The dredged specimens were chiefly extracted from tubes composed of plant
debris and sediment particles cemented to an organic lining and often the tubes were
found to run along the central cavities of hollow plant stems. Specimens from the
littoral zone had tubes encrusted with coarse sand grains and Halimeda discs.

DISTRIBUTION. Tropical Indo-west-Pacific to Japan.

Subfamily LYSARETINAE
Oenone fulgida (Savigny, 1818)

Aglaurides fulgida: Fauvel, 1953, 250, fig. 125. a-f.

Oenone fulgida: Crossland, 1924 : 85, fig. 106-111; Imajima & Hartman, 1964 : 267; Day,
1967 : 426, fig. 17.14. a-g.

HABITAT. Boring in coral (Porites).
RECORDS. Tetel Is. - 2 ; Yandina - i.
DISTRIBUTION. Circumtropical.

Subfamily LUMBRINERINAE
Lumbrineris latreilli Audouin & Milne-Edwards, 1834

Lumbriconereis latreilli: Fauvel, 1923 : 431, fig. 171. m-r; 1953 : 266, fig. 134. m-r.
Lumbrineris latreilli: Day, 1967 : 438, fig. 17.16. p-t.

HABITAT. Under coral boulders in silty sand; mud at 18 m depth.

164 P. E. GIBBS

RECORDS. Tetel Is. - I ; Komimbo Bay - i ; Graham Pt. - 2 ; Maraunibina Is. - I ;
ML 188 - i.

DISTRIBUTION. Cosmopolitan.

Lumbrineris papillifera (Fauvel, 1919)

Lumbriconereis papillifera Fauvel, igiga, : 395, pi. 15, figs 9-16.
Lumbrineris papillifera: Day, 1967 : 442, fig. 17.17. p-s.

HABITAT. Silty sand at 9 m depth.
RECORD. ML 56-1.

NOTES. The specimen is incomplete posteriorly but middle segments having the
characteristic ventral papillae are present and other details correspond to the species
description. This is the first record of the species from the Pacific region.

DISTRIBUTION. Tropical East Africa.

Lumbrineris sphaerocephala (Schmarda, 1861)

Notocirrus sphaerocephalus Schmarda, 1861 : 116, with text-figs.

Lumbriconereis sphaerocephala: Ehlers, 1904 : 33, pi. 5, figs 3-11; Fauvel, 1953 : 267, fig. 135. c-f.

HABITAT. Clean sand at LWM.
RECORD. Komimbo Bay - i.
DISTRIBUTION. Indo-west-Pacific.

Subfamily ARABELLINAE
Arabella tricolor (Montagu, 1804)

Arabella iricolor: Fauvel, 1923 : 438, fig. 175. a-h; 1953 : 2 74 ^g- I 4- a ~^'> Day, 1967 : 446,
fig. 17.18. i-m.

HABITAT. Under boulders in muddy silt, sand and shell gravel at MTL.
RECORDS. Tetel Is. - 3; Maraunibina Is. - 2; Graham Pt. - 6.
DISTRIBUTION. Cosmopolitan.

Arabella mutatis (Chamberlin, 1919)

Arabella mutans: Fauvel, 1953 : 275, figs 140. i-1, 143. g-i; Day, 1967 : 446, fig. 17.18. f-h.
HABITAT. Amongst Phyllochaetopterus socialis tubes at LWM.
RECORDS. Batuona Is. - 5.
DISTRIBUTION. North Atlantic; East Africa; Easter Is.; eastern Pacific.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 165

Drilonereis major Grassland, 1924

Drilonereis major Crossland, 1924 : 57, figs 73-79; Fauvel, 1953 : 277, fig. 143. k-1.

HABITAT. Boring (?) in coral on reef platform at LWM.
RECORDS. Maramasike Passage - 2.

NOTES. The prostomium is semi-circular and flattened. The maxillary formula
is MX. I = i + i ; MX. II = 6 + 6; MX. Ill = (3 4) + (3 4); MX. IV = (2 3)
+ (2 3) : the teeth of MX. Ill and MX. IV are very irregular in size and difficult
to count, as noted by Fauvel (1932). This is a new record for the Pacific region.

DISTRIBUTION. Gulf of Suez; Zanzibar; Bay of Bengal.

Subfamily DORVEILLEINAE
Dorvillea sp.

HABITAT. Coarse sand at MTL.
RECORD. Komimbo Bay - i.

NOTES. The specimen is 7 mm long (incomplete). Its antennae have eight
annuli and the dorsal cirri, commencing on setiger 2, are mounted on long cirrophores.
Forked setae are present from the first setiger and have unequal prongs. It is
close to D. incertus (Schmarda) but differs from this species in having forked setae
from the first, not the second, setiger.

Family ORBINIIDAE
Haploscoloplos bifurcatus Hartman, 1957

Haploscoloplos bifurcatus Hartman, 1957 : 2 77-

HABITAT. Mud and silty sand, 2-13 m depth.
RECORDS. ML 68 - 16 ; ML 69 - i ; ML 228 - 2.

NOTES. The thorax is composed of 17 to 20 setigers. Branchiae commence on
setigers 8, 9 or 10 as small triangular lobes, increasing in size on the posterior thoracic
segments. Characteristically, the post-setal lobes of the neuropodia in the posterior
half of the thorax (from setiger 12) are divided. Abdominal parapodia lack inter-
ramal cirri.

DISTRIBUTION. South and south-east Australia.

166 P. E. GIBBS

Naineris laevigata (Grube, 1855)

Naineris laevigata: Fauvel, 1927 : 22, fig. 7. a-1; 1953 : 310, fig. 163. a-1; Hartman, 1957 : 2 97.
pi. 35, figs 1-8; Day, 1967 : 539, fig. 32.2. a-f.

HABITAT. Medium to coarse sand, MTL to LWM.
RECORDS. Haroro - i ; Graham Pt. - 2.
DISTRIBUTION. Cosmopolitan in warm waters.

Family SPIONIDAE
Dispio maroroi n. sp.

(Fig. ii. A-D)

DESCRIPTION. All four specimens are incomplete. The holotype is 18 mm in
length with 46 segments and the largest of the paratypes is 45 mm long for 86 seg-
ments. The diameter of the body is about 2 mm.

The prostomium is pointed anteriorly and extends back as an occipital ridge to
the first segment where it forms a small occipital tentacle (fig. n. A). On either
side of this ridge there are conspicuous nuchal slits which are concealed at their
posterior ends by dorsal extensions of the peristomium. Two small eyes, not
visible from the dorsal aspect, are situated laterally on the prostomium. The palps
are missing on all specimens.

The parapodia of the first two segments differ from those of subsequent segments
in that the post-setal lobes of the notopodia are expanded, with 6 or 7 digitiform
processes along their free edges (fig. n. B). The lobes of subsequent notopodia have
smooth outlines. The post-setal neuropodial lobes of the first few segments are
rounded, becoming flatter in shape in later segments. Pre-setal lobes are low and
rounded.

Branchiae are present on all segments from setiger i. They are fused with the
post-setal notopodial lobes for most of their length and are slightly longer than the
lobes. Commencing between setigers 18 to 20, accessory branchiae appear on the
posterior side of the notopodia. Each is a palmate group of 6 to 8 lobes, each lobe
containing a vascular loop (fig. 11. c).

Apart from the first setiger, which has a bundle of long fine capillaries which
projects forward over the head region, the notosetae are broad- winged and narrow-
winged capillaries, the two types being of similar length. The neurosetae of the
anterior segments are similar to the notosetae except that the narrow-winged
capillaries are longer than the broad-winged. Between setigers 24 to 28, the broad-
winged capillaries are replaced by very finely pointed setae and 4 or 5 unidentate
hooded hooks appear (fig. n. D). In all neuropodia, there is an inferior group of
4 or 5 sabre-like setae.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS

167

D. mar or oi is close to the type species D. uncinata Hartman (1951) from Florida
but differs from it in that the post-setal notopodial lobes of setigers I and 2 are much
enlarged whilst that of setiger 3 is unmodified. A further difference is that the
accessory branchiae start between setigers 18 to 20 not between setigers 24 to 28 and
also there are minor variations in the form of the head region. In other details the
specimens from the Solomon Islands resemble those from Florida. The central
American species D. remanei and D. schusterae, described by Friedrich (1956), both
differ in having digitate neuropodial, as well as notopodial, lobes in the anterior
segments. In the fourth species, D. magna (Day, 1955) the accessory branchiae
take the form of minute lamellae, not lobes.

HABITAT. Clean sand at LWM.

RECORDS. Komimbo Bay - i (holotype) ; Graham Pt. - 3.
British Museum (Natural History) Registration No. Holotype 1970-53

Paratypes 1970-54

FIG. n. Dispio maroroi n.sp. (A) Dorsal view of head region. (B-C) Posterior views of
parapodia from setiger i (setae omitted) and setiger 40. (D) Neuropodial hook.

Laonice cirrata: Fauvel, 1927
fig. 18.6. h-k.

Laonice cirrata (Sars, 1851)
: 38, fig. 12. a-e; 1953 : 315, i

HABITAT. Mud at n m depth.
RECORD. ML 195 - i.
DISTRIBUTION. Cosmopolitan.

j. 165. a-e; Day, 1967 : 480,

168 P. E. GIBBS

Malacoceros indicus (Fauvel, 1928)

Scolelepis indica Fauvel, 1928 : 93, fig. 2. g-m; 1953 : 313, fig. 165. g-m.

Malacoceros indicus: Pettibone, I9&3a : 99 (synonymy); Day, 1967 : 477, fig. 18.5. p-u.

HABITAT. Silty mud and sand about MTL.
RECORDS. Nggela Is. - 2 ; Tetel Is. - i ; Fintry Pt. - 9.

NOTES. Specimens have 10 to 12 bidentate hooks in the neuropodia; in specimens
from India there are less than 6.

DISTRIBUTION. Indo-west-Pacific.

Nerinides sp. cf. gilchristi Day, 1961

Nerinides gilchristi Day, 1961 : 491, fig. 5. a-d; 1967 : 485, fig. 18.7. i-1.
HABITAT. Silty sand (Thalassia flat).
RECORD. Komimbo Bay - i.

NOTES. The specimen is in two parts - an anterior fragment of 15 setigers and a
middle body section of 13 setigers. In the latter, the branchiae are flag-shaped,
corresponding to those of N. gilchristi. However there are 7 or 8 hooded hooks per
neuropodium, each with a single tooth above the main fang, not 10 to 12 hooks having
three teeth above the main fang, as in N. gilchristi.

Polydorella novaegeorgiae n. sp.

(Fig. 12. A-F)

DESCRIPTION. All five specimens are incomplete. The largest fragment, chosen
for the holotype, measures 15 mm for 50 setigers and has a width of i-o to 1-5 mm.
The colour is whitish except for a brownish pigmentation on the peristomium and
anterior segments.

The prostomium has a rounded anterior margin and extends back as a low ridge
to the middle of setiger 2. On its dorsal surface there are two pairs of eyes and a
small occipital papilla (fig. 12. A). The broad peristomium carries a pair of stout
palps which reach to about setiger 15.

On the anterior segments, the notopodial lobes gradually increase in size and are
largest on setiger 4. The neuropodial lobes of setigers i to 6 are similar in size and
are semi-circular in outline; from setiger 7 they are small and inconspicuous.
Branchiae commence on setiger 6 and continue to setigers 15 to 18 (i.e. there are 10
to 13 pairs). These are cirriform in shape, the longest being equal to the body width.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS

169

Both notosetae and neurosetae in the anterior segments, with the exception of the
notosetae of setiger 4, are winged capillaries. The notopodial bundles are composed
of two rows of curved capillaries which have narrow blades on one side (fig. 12. c),
plus a superior group of similar form but much longer. The neurosetae are narrow-
winged capillaries, each finely tapering to a sharp point.

As is characteristic of the genus, the notopodium of setiger 4 carries modified
setae. Two types are present in a double row that follows a wide arc (fig. 12. B) ;
those of the anterior row are strongly angled at their distal end with a broad blade
showing fine striations (fig. 12. D) while those of the posterior row are stout hooks
with bluntly tapering, slightly curved tips (fig. 12. E). There are between 20 and
25 pairs of the two types of modified setae, plus a superior group of about 15 narrow-
winged capillaries, in each of the notopodia of setiger 4. In the post-branchial
region the superior capillary group is replaced by a group of about 10 needle-like
setae. Bidentate hooded hooks (fig. 12. F) commence at setiger 7 and number
between 18 and 20 per neuropodium.

The type species Polydorella prolifera Augener was described from Sharks Bay,
Western Australia, (Augener, 1914) and has been recorded from India (Gravely,

FIG. 12. Polydorella novaegeorgiae n.sp. (A) Dorsal view of anterior region. (B) Lateral
view of setiger 4. (c) Notopodial capillary. (D-E) Modified notopodial setae from
setiger 4. (F) Neuropodial hook.

iyo P. E. GIBBS

1927; Fauvel, 1930). It differs from P. novaegeorgiae in many respects, particularly
in the form of the prostomium, the structure of the modified setae of setiger 4 and
in the number of branchiae. A further point of difference is that the specimens
from Marovo Lagoon do not show any sign of reproduction by scissiparity, a feature
observed in P. prolifera.

HABITAT. Mud at n m depth.

RECORDS. ML 39-5.

British Museum (Natural History) Registration No. Holotype 1970-55

Paratypes 1970-56

Prionospio cirri/era Wire"n, 1883

Prionospio cirrifera : Fauvel, 1927 : 62, fig. 2i.k-n; 1953 : 324, fig. 164. k-m; Day, 1967 : 486,
fig. 18.8. a-d.

HABITAT. Mud at 22 m depth.

RECORD. ML 29-1.

NOTES. Although P. cirrifera has not been recorded previously from the west
Pacific region the single specimen agrees in detail. It has 12 pairs of branchiae and
the hooded crotchets are multidentate, corresponding to the northern form (cf. Day,
1967).

DISTRIBUTION. North Atlantic; North Pacific; India; South Africa.

Prionospio ehlersi Fauvel, 1928
Prionospio ehlersi Fauvel, ig28a :'io, fig. i. a-r; Day, 1967 : 490, fig. 18.9. d-f.

HABITAT. Mud and sand, 2-22 m depth.

RECORDS. ML 37 - 2 ; ML 41 - 4 ; ML 134 - 1 ; ML 156- 8 ; ML 191 - 1 ; ML 194 -2 ;
ML 195-1; ML 218 -6.

NOTE. This is the first record of P. ehlersi from the Pacific region.

DISTRIBUTION. Morocco; East Africa.

Prionospio malmgreni Claparede, 1870

Prionospio malmgreni: Fauvel, 1927 : 61, fig. 21. a-e; Hartman & Imajima, 1964 : 285; Day,
1967 : 492, fig. 18.9. a-c.

HABITAT. Silty sand at 2 m depth.
RECORDS. ML 196 - 2.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 171

NOTES. Both specimens are small, about 10 mm long. Across setiger 7 there is
a membraneous ridge and this feature separates P. malmgreni from the closely allied
species P. steenstrupi Malmgren.

DISTRIBUTION. Atlantic; Mediterranean; South Africa; South-west Australia;
Japan; southern California.

Prionospio pinnata Ehlers, 1901

Prionospio pinnata : Fauvel, 1953 : 323, fig. 174. e; Imajima & Hartman, 1964 : 286; Day, 1967 :
488, fig. 18.8. i-1.

HABITAT. Mud and silty sand, 2-26 m depth

RECORDS. ML 37-10; ML 40-5; ML 41 -27;. ML no -4; ML 156- 2; ML
157 - 2 ; ML 191 - i ; ML 194 - 2 ; ML 195 - i.

DISTRIBUTION. Cosmopolitan.

Prionospio steenstrupi malayensis Caullery, 1914

Prionospio steenstrupi malayensis: Caullery, 1944 : 14, fig. 8. a-f.

HABITAT. A wide variety of deposits from mud to coarse sand, 2-24 m depth.

RECORDS. ML 55 - 2 ; ML 68 - i ; ML 188 - i ; ML 192 - i ; ML 196 - 5 ; ML 204 -
i; ML 230- i.

NOTES. Caullery recognized the subspecies malayensis on the grounds of geo-
graphical separation from the stem species which was then (1914) known only from
the North Atlantic. In the Solomon Islands specimens there are low membraneous
ridges across the dorsum from setiger 12 to setiger 16 or 17. These ridges are not
present in the stem species and thus this feature distinguishes the two forms.

DISTRIBUTION. East Indies.

Prionospio tetelensis n. sp.

(Fig. 13. A-D)

DESCRIPTION. All specimens are incomplete. The holotype from ML 188 is
14 mm long for 42 segments and has a diameter of 2 to 3 mm. One of the paratypes
(from ML 68) is a mature female containing oocytes which are turquoise blue in
colour.

The prostomium is rounded anteriorly and extends back to setiger 3 as an elevated
keel between peristomial folds (fig. 13. B). There are three pairs of prostomial

I 7 2

P. E. GIBBS

eyes, two pairs of which are small and situated to the anterior of a larger, transverse
pair. On either side of the prostomium, the peristomium is extended to form lateral
wing-like expansions.

The first setiger is small and lacks notosetae. The parapodial lobes of the anterior
segments are large and ear-shaped, the pre-setal ones being smaller than the post-
setal (fig. 13. c). These lobes gradually decrease in size, becoming flatter and less
rounded in the middle body. From setiger 6 to between setigers 20 to 26 the post-
setal notopodial lobes are joined across the dorsum by membraneous ridges. In
mature specimens there are no genital pockets between the anterior neuropodia.

Branchiae are present on setigers 2 to 5 and all are pinnate (fig. 13. A). Each has
a short stem and numerous pinnules arising from the lateral and posterior surfaces
of the main axis. The size of the branchiae decreases from the first to the fourth
pair.

Both notosetae and neurosetae in the anterior segments are narrow-winged capil-
laries. Commencing between setigers 20 and 22, about 12 hooks appear in the
neuropodia, accompanied by very fine pointed setae and also 2 or 3 sabre-like setae.
Hooks do not appear in the notopodia until about setiger 50. Each hook is hooded
with a series of five teeth above the main fang (fig. 13. D).

FIG. 13. Prionospio tetelensis n.sp. (A-B) Lateral and dorsal views of anterior
region, (c) Parapodium of setiger 15. (D) Neuropodial hook.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 173

Of the many species of the genus Prionospio now known, three are similar to P.
tetelensis in possessing four pairs of pinnate branchiae on setigers 2 to 5. These are
P. treadwelli Hartman, P. tennis (Verrill) - both from the Atlantic coast of North
America - and P. peruana Hartmann-Schroder from Peru. P. tetelensis may be
distinguished from all three species on the basis of a combination of characters,
namely, the prostomial keel between peristomial folds extending to setiger 3, the
lateral wing-like expansions of the peristomium, the membraneous ridges across the
dorsum of setigers 6 to 20 and the neuropodial hooks being present from setiger 20.

HABITAT. Mud from LWM to 18 m depth.

RECORDS. Tetel Is. - i ; ML 68 - i ; ML 188 - i (holotype) ; ML 195 - i ; ML
228 - 1.

British Museum (Natural History) Registration No. Holotype 1970-57

Paratypes 1970-58-61

Pseudopolydora corallicola Woodwick, 1964

Pseudopolydora corallicola Woodwick, 1964 : 151, fig. 2. 9-12.

HABITAT. Coralline mud at LWM ; silty sand at 2 m depth.
RECORDS. Matiu Is. - i ; ML 196 - i.

NOTES. Both specimens are small (under 5 mm long) and are damaged. Those
characters that can be checked, such as the prostomial ridge extending to setiger 6,
the modified setae of setiger 5 and the peculiar fenestrated neuropodial hooks, all
correspond.

DISTRIBUTION. Marshall Islands.

Pseudopolydora sp.

HABITAT. In brackish water (5-6 % ) - boring in sponge.

RECORDS. Lagoon at the mouth of the Lunga River at Lunga Point - numerous
specimens.

NOTES. These specimens are up to 5 mm in length. The modified setae of
setiger 5 include an anterior row of broad-winged, sharply tapering setae and a
posterior row of stout hooks with recurved tips: there are 6 or 7 pairs of the two
types. Branchiae commence on setiger 6 and extend to setigers 16 or 17 (n or 12
pairs). Neuropodial hooks are present from setiger 8. In the posterior notosetae
there are one or two stout, curved hooks which arch over the dorsum.

Mr. W. J. Light has examined this series and, in a personal communication, he

174 P. E. GIBBS

has pointed out that certain characters, in particular the form of the modified setae
on setiger 5 and also the presence of specialized posterior notosetae, indicate that
these specimens represent a new species of Pseudopolydora. He has kindly consented
to publish a description in a separate paper.

Scolelepis squamata mendanai n. subsp.

DESCRIPTION. The holotype is a complete specimen measuring 50 mm in length
for about 200 segments. The diameter of the body is between i-o to 1-5 mm. The
palps are very small and slender extending only to setiger 3 or 4. There are four
eyes set in a transverse row across the prostomium. Branchiae commence on setiger
2 and are fused to the notopodial lamellae for about one-third of their length.
Bidentate hooks number 9 or 10 in the neuropodia from about setiger 45, and I to 3
in the notopodia starting between setigers 90 to 100. The post-setal neuropodial
lobes are ear-shaped in anterior segments but become bilobed between setigers 35 to
45 and are distinctly divided in the middle segments.

The subspecies mendanai is distinct from the stem species (Fauvel, 1927 : 36,
fig. ii. g-n, as Nerine cirratulus) in the following characters: (i) the palps are very
small reaching only to setiger 3 (not setiger 24), (ii) neuropodial hooks are present
from setiger 45 (not setiger 40) and (iii) notopodial hooks are present from about
setiger 90 (not setiger 60). The subspecies saipanensis (Hartman, 19540 : 230, fig.
2. a-e, as Nerine cirratulus saipanensis) differs in that the hooks appear in the neuro-
podia from setigers 29 to 31 and in the notopodia from setiger 41.

HABITAT. Silty sand, MTL to LWM.

RECORDS. Graham Pt. - 5 ; Fintry Pt. - 6 (including holotype).
British Museum (Natural History) Registration No. Holotype 1970-62

Paratypes 1970-63-65

Spio filicornis (Muller, 1776)

Spiofilicornis: Fauvel, 1927 : 43, fig. 15. a-g; Imajima & Hartman, 1964 : 289; Day, 1967 : 481,
fig. 18.6. l-o.

HABITAT. Coralline mud at LWM.

RECORDS. Matiu Is. - 8.

DISTRIBUTION. North Atlantic ; South Africa ; North Pacific.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 175

Family MAGELONIDAE
M agelona japonica Okuda, 1937

Magelona japonica Okuda, 1937 : 247, figs 23-24; Imajima & Hartman, 1964 : 290; Tampi &
Rangarajan, 1964 : 113, figs 44-49.

HABITAT. Coarse sand below LWM.

RECORDS. Tetel Is. - 2.

DISTRIBUTION. Bay of Bengal; North Pacific.

Magelona sp.

HABITAT. Silty sand at 2 m depth.
RECORD. ML 283 - i.

NOTES. The specimen is an anterior fragment of 20 segments. The prostomium
is elongated with a rounded anterior margin lacking frontal horns. Anterior segments
have large notopodial and small neuropodial lamellae. The setae of segment 9 are
unspecialized. From setiger 10 the parapodial lamellae are rhomboidal. The
abdominal hooded hooks are bidentate with a single tooth above the main fang.

The specimen is close to M. capensis Day, differing in the shape of the abdominal
lamellae being rhomboidal not oval and in the hooks being bidentate not tridentate.

Family TROCHOCHAETIDAE
Poecilochaetus serpens honiarae n. subsp.

DESCRIPTION. All of the specimens are incomplete; the largest specimen (the
holotype) is 20 mm long with 44 segments and has a body width of 2-0 mm. The
characters are those of the stem species (Allen, 1904 : 79, pi. 7-12, text-fig, i) except
that the aristate spines, with knobbed ends and hairy terminal aristae (Allen, 1904 :
pi. 9, fig. 18) commence between setigers 20 to 25 and thus are not confined to the
posterior region beyond setiger 80, as in the stem species.

HABITAT. Silty and coarse sand, MTL to LWM.

RECORDS. Tetel Is. - 3 (including holotype) ; Komimbo Bay - 2.
British Museum (Natural History) Registration No. Holotype 1970-66

Paratypes 1970-67-69

NOTES. The Solomon Islands specimens have a reddish, chitinous, triangular
plate on the dorsum of setiger 9 (a similar, but diamond-shaped, structure has been

176 P. E. GIBBS

noted in P. johnsoni by Hartman, 1939). Lateral sense organs are missing from
setigers 6 to 9 and the branchiae, composed of two filaments on the posterior sides
of the parapodia, are present from setigers 17 or 18.

An examination of P. serpens individuals from the type locality at Plymouth
reveals that Allen overlooked the dorsal plate on setiger 9 and also the absence of
lateral sense organs on setigers 6 to 9. Allen states that there are two pairs of
branchial filaments from setiger 21 but in the several specimens which have been
examined there is only one pair, although a third filament sometimes occurs after
setiger 32.

Thus adult specimens of the stem species P. serpens from the type locality and
adults of the subspecies honiarae from the Solomon Islands are very similar, the
distinguishing feature being the presence of aristate spines in the middle body seg-
ments of the latter. This distinction is recognized as a minor one, deserving of sub-
specific separation only.

P. serpens is remarkable in its retention of the pelagic habit until a late stage of
larval development. Thus it is not surprising that the existence of this species in the
Indo-west Pacific region was known from plankton records long before adult speci-
mens were discovered in the region. Pelagic stages have been described from Indo-
China (Fauvel, 1939) and India (Banse, 1959; Ganapati & Radhakrishna, 1958).
Only recently has an adult specimen been discovered in Indian waters (Achari, 1968).

Poecilochaetus tropicus Okuda, 1935

Poecilochaetus tropicus Okuda, 1935 : 289, figs 1-2; I937a : 294, figs 39-40.
HABITAT. Silty and coarse sand from MTL to LWM.
RECORDS. Tetel Is. - 5; Graham Pt. - i.

NOTES. P. tropicus was described from an incomplete specimen of 39 segments
taken in the Palau Islands in 1934. No further material appears to have been
recorded since that date.

All of the Solomon Islands specimens are incomplete. The most complete
individual is immature, measuring 26 mm long for 70 setigers, but the largest specimen
appears mature and has a length of 30 mm for 50 setigers. The original description
can be supplemented with the following details - (i) the two branchial filaments
situated on the posterior sides of the parapodia are not present in small specimens
but are conspicuous in mature individuals; (ii) on the dorsal surface of setiger 9,
there is a small, yellowish, semi-lunar-shaped plate (a similar structure to that in
P. serpens)', (iii) the flask-shaped parapoch'al lobes of setigers 7 to 13 contain an
iridescent, light blue material in mature specimens, (iv) the spines with hairy
terminal filaments (Okuda, 1935, fig. 2.e) are not replaced by the knobbed, aristate
type (as in P. serpens) in the region anterior to setiger 70 in the most complete
specimen available.

DISTRIBUTION. Palau Islands.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 177

Family CHAETOPTERIDAE
Chaetopterus variopedatus (Renier, 1804)

Chaetopterusvariopedatus: Fauvel, 1927 : 77, fig. 26. a-n; 1953 : 337, fig. 175. a-n; Day, 1967 :
529, fig. 22.2. a-g.

HABITAT. Attached to Acropora on wharf piles; in sand at 2 m depth.
RECORDS. Yandina - 5 ; ML 118-2.

NOTE. All of the specimens are small, the largest being only 20 mm long.
DISTRIBUTION. Cosmopolitan.

Mesochaetopterus Sagittarius (Claparede, 1870)

Mesochaetopterus minutus Potts, 1914 : 963, pi. 2, fig. 4, pi. 3, figs 7-8, text-figs 4-5; Fauvel,

1953 : 342, fig- i? 8 - a; Da Y. J 96? : 53L fi g- 22.2. h-n.
Mesochaetopterus Sagittarius: Bhaud, 1969 : 325, fig. i. a-f.

HABITAT. A wide variety of deposits from coralline mud to coarse sand, MTL to
LWM ; in Halichondria at 5 m depth.

RECORDS. Tetel Is. - 25 ; Komimbo Bay - 22 ; Matiu Is. - i ; Maraunibina Is. - 2 ;
Graham Pt. - i ; Pigeon Is. - num. ; Fintry Pt. - 8 ; Yandina - num.

NOTES. The type specimens measure 25 mm long with a diameter of i mm
(Potts, 1914) but Monro (1931) records specimens from N.E. Australia that are twice
this size (50 x 2 mm) with tubes up to 20 cm in length. Some of the specimens
from Tetel Is. and Komimbo Bay, and all of those from Pigeon Is. and Yandina are
similar in size to the type material but the remaining specimens are larger and
approximate the size of the Barrier Reef specimens.

DISTRIBUTION. Mediterranean ; Indo-west-Pacific to Japan.

Phyllochaetopterus elioti Crossland, 1903

Phyllochaetopterus elioti Crossland, 1903 : 172, pi. 16, figs 1-4, 7-8, pi. 17, figs 10-13; Fauvel,
1953 : 34. fi g- 177- e - h : Da Y. 1967 : 525. fi g- 22.1. f-g.

HABITAT. Abundant in silty sand, MTL to LWM ; coarse sand at 4 m depth.
RECORDS. Tetel Is. - num. ; Fintry Pt. - num. ; ML 295 - i.
DISTRIBUTION. South and East Africa; India.

178 P. E. GIBBS

Phyllochaetopterus herdmani (Hornell, 1903)

Phyllochaetopterus herdmani: Willey, 1905; 292, pi. 5, figs 127-132; Fauvel, 1953 : 342, fig. 177.
i-m; Day, 1967 : 524, fig. 22.1. a-e.

HABITAT. Silty sand at MTL.
RECORD. Tetel Is. - i.

NOTE. The single specimen is rather damaged and in a poor state of preservation
but nevertheless corresponds to the species description. It is anterior fragment of
31 setigers comprising n anterior, 2 middle and 18 abdominal segments. On setiger
4 there are 9 modified setae in each of the notopodia. This is a first record for the
Pacific region.

DISTRIBUTION. East Africa; Ceylon.

Phyllochaetopterus socialis Claparede, 1870

Phyllochaetopterus socialis : Fauvel, 1927 : 84, fig. 30. a-1; 1953 : 339, fig. 176. a-1; Day, 1967 :
525, fig. 22.1. h-r.

HABITAT. Very abundant on moderately exposed reef platforms, forming dense
colonies towards LWM (see Gibbs, 1969, fig. 136); single specimens embedded in
Milkpora and the sponge Neofolitispa dianchora (de Laubenfels) from 5 m depth.

RECORDS. Matiu Is., Cape Esperance and Batuona Is. - numerous specimens;
Maramasike Pg. - 3; Kokomtambu Is. - i ; Yandina - i.

DISTRIBUTION. Mediterranean; Atlantic; Indo-Pacific.

Spiochaetopterus costarum costarum (Claparede, 1868)

Telepsavus vitrarius Ehlers, 1908 : 114, pi. 15, figs 1-8.
Spiochaetopterus vitrarius: Day, 1967 : 528, fig. 22.1. u-v.
Spiochaetopterus costarum costarum: Gitay, 1969 : 14 (synonymy).

HABITAT. Silty sand (Thalassia flat) at about MTL ; coarse sand at LWM.
RECORDS. Tetel Is. - 6 ; Komimbo Bay - 14.

NOTES. The specimens have a rounded prostomium with a pair of lateral eyes.
The anterior region is composed of 9 setigers, the notopodia of setiger 4 bearing single
modified setae with swollen, triangular or heart-shaped tips. On the ventral
surface of setigers 7 to 9 the tissue is distinctly glandular, being brownish-black in
colour on setiger 7 but whitish on setigers 8 and 9. On setiger 2 there is usually a
thin line of brownish pigment across the ventrum. Up to 19 middle and 25 ab-

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 179

dominal segments have been noted in several specimens but most of the material is
poorly preserved. The largest specimens measure about 25 mm in length, their
tubes being up to 60 mm long and 0-5 to i-o mm in diameter. The latter are trans-
parent and highly wrinkled.

Gitay (1969) has recognized three subspecies of 5. costarum in the Pacific region,
namely pottsi, monroi and okudai. The specimens from the Solomon Islands differ
from all three in having fewer middle segments (i.e. 19 compared to 30-90, about 50
and about 67 respectively). Furthermore their small size distinguishes them from
pottsi and the presence of eyes separates them from monroi and okudai. However
these specimens appear to be identical with T. vitrarius Ehlers, known from the
eastern Atlantic and Natal, which Gitay considers to be a synonym of S. costarum
costarum.

DISTRIBUTION. Eastern Atlantic; Mediterranean; Indian Ocean.

Family GIRRATULIDAE
Cirriformia filigera (Delle Chiaje, 1825)

Audouinia filigera: Fauvel, 1927 : 92, fig. 32. h-m; 1953 : 331, fig. 173. h-1.
Cirriformia filigera Day, 1967 : 518, fig. 20.4. p-q.

HABITAT. In Acropora rubble and under coral boulders on reef platforms; in
crevices and vacated borings in Porites, Acropora and beachrock.

RECORDS. Kokomtambu Is. - i ; Tetel Is. - 3 ; Komimbo Bay - 3 ; Matiu Is. - i ;
Paleki Is. - 2; Lauvie Is. - 2.
DISTRIBUTION. Atlantic; Mediterranean; Indian Ocean.

Cirriformia punctata (Grube, 1859)

Audouinia semicincta: Fauvel, 1953 : 330 fig. 174. c; Okuda, 1937 : 2 96, fig. 41. a-c.
Cirriformia punctata: Day, 1967: 517, fig. 20.4. j-m.

HABITAT. In sedimented crevices on reef platform.
RECORDS. Komimbo Bay - i ; Batuona Is. - 54.
DISTRIBUTION. Circumtropical.

Dodecaceria fistulicola Ehlers, 1901

Dodecaceria fistulicola : Fauvel, 1953 : 335, figs 169. h-i, 174. a-b; Okuda, 1937 : 2 98, fig. 42. a-f.
HABITAT. Crevices in reef platform.
RECORDS. Komimbo Bay - i ; Matiu Is. - i.

180 P. E. GIBBS

NOTES. A complete specimen has 68 segments and is about 15 mm long. There
are 4 or 5 pairs of branchiae which are shorter than the palps. The first 7 or 8
setigers carry capillary setae which are replaced in later segments by 5 or 6 hooks in
both rami.

DISTRIBUTION. Indo-west-Pacific ; southern South America.

Dodecaceria laddi Hartman, 1954

Dodecaceria laddi Hartman, 1954 : 638, n 8 s I 7^- c > J 77- d-h; Day, 1967 : 502, fig. 20.1. g-i.

HABITAT. Boring (?) in Forties.
RECORD. Kokomtambu Is. - i.
DISTRIBUTION. Marshall Islands; South Africa.

Tharyx sp.

HABITAT. Silty sand at 36 m depth.
RECORD. Off Fintry Pt. - i.

NOTES. The specimen is incomplete with 26 segments and measures 3-5 mm long.
The diameter is about 0-4 mm. The palps have been lost but scars indicate their
position. The setae are entirely capillary. It is impossible to identify this fragment
with any certainty but possibly it is T. multifilis Moore, which is known from Madras
(Fauvel, 1953).

Family GOSSURIDAE
Cossura coasta Kitamori, 1960

Cossura coasta Kitamori, 1960 : 1082, fig. i. a-f; Imajima & Hartman, 1964 : 299; Day, 1967 :
581, fig. 26.1. a-d.

HABITAT. Mud at 22 m depth.

RECORD. ML 37 - 2.

NOTES. Both specimens are incomplete; the larger measures 10 mm for 60
segments and has a diameter of 0-5 mm. They differ from the species description
only in respect of the shorter capillary setae which, instead of being serrated, are
smooth, and also taper quite abruptly about halfway along their length to a very
slender filament.

DISTRIBUTION. Southern Africa; Japan.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 181

Family FLABELLIGERIDAE
Diplocirrus glaucus orientalis n. subsp.

DESCRIPTION. The specimen chosen for the holotype (from ML 194) is 18 mm long
for 25 segments and has a maximum diameter of about 2-0 mm in the anterior body
region. A complete paratype (from ML 55) is 20 mm in length for 40 segments.
The characters are those of the stem species Diplocirrus glaucus (Malmgren), (Fauvel,
1927 : 120, fig. 43. a-d) but is distinguished by the presence of conspicuous orange
coloured, globular papillae situated on the ventral surface below each neuropodium
from setiger 4 to setigers 14 or 16.

HABITAT. Mud and silty sand, n to 24 m depth.

RECORDS. ML 55 - i ; ML 188 - 3 ; ML 194 - i (holotype) ; ML 218 - i ;
ML 230 - 1.

British Museum (Natural History) Registration No. Holotype 1970-70

Paratypes 1970-71-74

NOTES. The stem species is widely distributed in the North Atlantic region and,
although Fauvel (1932) refers a specimen dredged in the Mergui Archipelago from
7 m (4 fm) to this species 'with great hesitation', no certain record of this species
being taken in the Indian Ocean exists.

Specimens of the stem species from Plymouth do not possess conspicuous papillae
below the neuropodia of the anterior segments. The presence of these papillae thus
distinguishes the subspecies orientalis.

Family SGALIBREGMIDAE
Hyboscolex longiseta Schmarda, 1861

Hyboscolex longiseta Schmarda, 1861 : 54, pi. 27, fig. 211, with text-figs; Day, 1967 : 588, fig.
27.2. a-d.

HABITAT. Under coral boulders in muddy silt ; in beachrock.
RECORDS. Tetel Is. - i ; Maraunibina Is. - i ; Lauvie Is. - 2.
DISTRIBUTION. Southern and East Africa; New Zealand (Auckland Is.).

Scalibregma inflatum Rathke, 1843

Scalibregma inflatum: Fauvel, 1927 : 123, fig. 44. a-f; 1953 : 355, fig. 185. a-f; Day, 1967 : 590,
fig. 27.2. e-i.

HABITAT. Silty mud at 13 m depth.

RECORD. ML 69-1.

DISTRIBUTION. Cosmopolitan (but few records from the Indo-Pacific region).

i8a P. E. GIBBS

Family OPHELIIDAE
Armandia lanceolata Willey, 1905

Armandia lanceolata Willey, 1905 : 288, pi. 5, fig. 120; Fauvel, 1953 : 358.

HABITAT. Acropora rubble at LWM; silt and sand, 5-24 m depth.

RECORDS. Tetel Is. -i; ML 56-1; ML 117 - i; ML 134 - i; ML 229 - i;
ML 230 - 1.

DISTRIBUTION. Indo-west-Pacific.

Armandia leptocirrus (Grube, 1878)

Ophelina (Armandia) leptocirrus Grube, 1878 : 194.

Armandia leptocirrus: Fauvel, 1953 : 358; Day, 1967 : 577, fig. 25.2. h.

HABITAT. Silt and coralline mud at LWM ; mud and sand, 2-18 m depth.

RECORDS. Komimbo Bay - i ; Matiu Is. - 7 ; ML 68 - 2 ; ML 97 - i ; ML 118 - i
ML 190 - 2 ; ML 192 - i ; ML 196 - 3 ; ML 283 - 4.

DISTRIBUTION. Tropical Indo-west-Pacific.

Armandia longicaudata (Caullery, 1944)

Ammotrypane longicaudata Caullery, 1944 : 44, fig. 35. a-c.
Armandia longicaudata: Day, 1967 : 577, fig. 25.2. a-c.

HABITAT. Silty sand at 24 m depth.

RECORD. ML 230 - i.

DISTRIBUTION. East Africa; East Indies.

Ophelia koloana n. sp.

(Fig. 14. A-B)

DESCRIPTION. The two type specimens measure 10 to 12 mm in length and have
a diameter between 1-0-1-5 mm. The paratype is a female approaching maturity,
containing coelomic oocytes.

The prostomium is sharply pointed. The anterior region is cylindrical with the
ventral groove commencing at setiger 8. The first 9 setigers are abranchiate,

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 183

branchiae being present from setiger 10 to setiger 27 (i.e. there are 18 pairs). Each
branchia is rather small and has a 'crimped' appearance (fig. 14. A). The branchi-
ferous region is followed by 4 abranchiate segments and then two achaetous annuli
(or pre-anal segments) . Thus, following the scheme of Tebble (1953) , the body formula
is ga + i8b ( ) + 4a + 2n = 33. From setiger 30 to the pygidium there is a pair
of prominent dorso-lateral ridges (fig. 14. B). The pygidium bears two stout ventral
papillae and 9 small dorsal ones. Neither nephridiopores nor branchial fenestrations
in the lateral body wall could be detected.

In terms of its affinities, 0. koloana has a similar body formula to 0. denticulata
Verrill and 0. capensis Kirkegaard. However in 0. denticulata, which is an Atlantic
species, the ventral groove starts at segment 10 and it has n to 18 dorsal anal
papillae (Fauvel, 1927 : 132, fig. 46, g-h, as 0. neglecta, fide Tebble, 1953). 0.
capensis from South Africa differs in that it has lateral swellings from segment 26 to
the pygidium (Kirkegaard, 1959 : 45, fig. 8) not dorso-lateral ridges from setiger 29.

HABITAT. Sand at 2 m depth.

RECORDS. ML 118-2.

British Museum (Natural History) Registration No. Holotype 1970-75

Paratype 1970-76

NOTE. Hitherto the genus Ophelia appears to have been recorded only once in
the tropical Indo-west-Pacific region, namely, a single specimen of 0. limacina from
Indo-China (Fauvel, 1939).

0-5mm 0-5mm

FIG. 14. Ophelia koloana n.sp. (A) Lateral view of segments 22 to 24.
(B) Dorso-lateral view of posterior region.

Polyophthalmus pictus (Dujardin, 1839)

Polyopht halmus pictus : Fauvel, 1927 : 137, fig. 48. l-o; 1953 : 3^> & 1 ^7- l-o; Day, 1967 : 579,
fig. 25.2. k-m.

HABITAT. Interstices of the sponge Halichondria at 5 m depth.
RECORDS. Yandina - 3.

184 P. E. GIBBS

NOTES. All three specimens appear to be juvenile in that they measure only
3 to 7 mm in length. Pillai (1965) found this species in a similar habitat, namely,
within sponges and other encrusting organisms on bamboo oyster spat collectors.

DISTRIBUTION. Cosmopolitan.

Family STERNASPIDAE
Sternaspis scutata (Renier, 1807)

Sternaspis scutata: Fauvel, 1927 : 216, fig. 76. a-g; 1953 : 401, fig. 210. a-g; Day, 1967 : 648,
fig. 31.1. a-d.

HABITAT. Mud and silty sand, 18-24 m depth.
RECORDS. ML 218 - 6 ; ML 230 - 3.
DISTRIBUTION. Cosmopolitan.

Family CAPITELLIDAE
Capitellethus dispar (Ehlers, 1907)

Capitellethus dispar: Fauvel, 1953 : 37*^ Rullier, 1965 : 194.

HABITAT. Silty sand above LWM.
RECORDS. Komimbo Bay - 3.

NOTES. The material is fragmentary and the largest specimen measures 18 mm
long for 60 segments. The'peristomium is achaetous and there are n thoracic
setigers with capillary setae in both rami, which are replaced by hooks from setiger
12. No branchiae could be seen. In all three specimens the epidermal cells of
setigers n and 12 (the last thoracic and first abdominal) contain a dark pigment.

DISTRIBUTION. India; West Pacific.

Capitobranchus sp.

HABITAT. Coarse sand at LWM.
RECORD. Tetel Is. - i.

NOTES. The single specimen is incomplete, measuring 20 mm for 20 segments,
and has a diameter of about 2 mm. The peristomium is biannulate, segments 2 to
4 are triannulate, segments 5 to 8 are quadriannulate and the remaining segments are

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 185

biannulate. Segments 2 and 3 are achaetous but may have lost their setae. Seg-
ments 4 to 12 have capillaries in both rami which are replaced by hooks in the
neuropodia from segment 13.

In having at least 19 thoracic segments and hooks from setiger 13 (assuming
segments 2 and 3 to have been setigerous), the specimen falls within the broad
definition of the genus Capitobranchus given by Day (1962, p. 651) as follows:
"Thorax with an achaetous peristome followed by about 19 setigerous segments
most of which bear capillary setae in both rami, but hooded hooks are present in the
last few neuropodia." The genus is monospecinc based on C. macgregori from
Madagascar which has 19 thoracic setigers, the last four of which bear neuropodial
hooks. Due to its incomplete nature, the specimen from Tetel Island must remain
indeterminable as to species until further material becomes available.

Dasybranchus caducus (Grube, 1846)

Dasybranchus caducus: Fauvel, 1927 : 148, fig. 52. a-h; 1953, 365, fig. 190. a-h; Day, 1967 : 603,
fig. 28.3. e-h.

HABITAT. A wide variety of deposits, mud to coarse sand, particularly under coral
boulders on reef platforms, from above MTL to LWM ; mud, 9-22 m depth.

RECORDS. Tetel Is. - 56; Nggela Is. (Sandfly Pg.) - 30; Naro Bay - 7; Komimbo
Bay - 14; Matiu Is. - 16; Graham Pt. - 19; Fintry Pt. - 7; ML 29-2; ML 56 - I.

DISTRIBUTION. Atlantic; Mediterranean; Indo-Pacific.

Mastobranchus dollfusi Fauvel, 1936
Mastobranchus dollfusi Fauvel, 1936 : 81, fig. n. a-g; Kirkegaard, 1959 : 51.

HABITAT. Muddy sand at MTL; mud and silty sand, 2-22 m depth.

RECORDS. Nggela Is. - 4 ; ML 29 - i ; ML 68 - 5 ; ML 195 - i.

NOTES. Specimens have a diameter of 0-5 to i-o mm and the largest anterior
fragment is 30 mm long for 70 segments. The prostomium is conical with a pair of
pigmented eye-spots and is followed by an achaetous peristomium. The first setiger
has capillary notosetae only; setigers 2 to 10 have capillaries in both rami. The
number of thoracic setigers varies, as noted by both Fauvel and Kirkegaard for the
West African material. In the Solomon Islands specimens setiger n usually has
capillary notosetae and neuropodial hooks with the succeeding segments carrying
hooks in both rami. However two specimens have capillary notosetae on setiger 12,
one of which also has capillary neurosetae on setiger n. A fragment of the posterior
region possesses retractile branchiae situated posteriorly to the notopodia of the
abdominal segments, and the pygidium has four short anal cirri.

Despite the wide geographical separation of the two localities, the Solomon Islands
specimens appear to be identical to those from West Africa in all details.

DISTRIBUTION. West Africa.

186 P. E. GIBBS

Mastobranchus trinchesii Eisig, 1887

Mastobranchus trinchesii: Fauvel, 1927 : 152, fig. 54. a-i; Kirkegaard, 1959 : 52.

HABITAT. Silty sand, MTL to LWM.

RECORDS. Tetel Is. - i ; Komimbo Bay - 4; Graham Pt. - 27.

NOTES. Although M. trinchesii is a species hitherto known only from the
Mediterranean and West Africa, the Solomon Islands specimens appear to be identical.
Briefly, the latter specimens have an achaetous peristomium, followed by n thoracic
setigers with capillary setae in both rami, except the first which has notosetae only.
From setiger 12 the capillaries are replaced by hooks in the neuropodia, but in the
notopodia capillaries persist with the hooks until about setiger 90. Retractile
cirriform branchiae are present in the posterior segments and the pygidium carries
four anal cirri.

DISTRIBUTION. Mediterranean; West Africa.

Mediomastus sp. cf. capensis Day, 1961

Mediomastus capensis Day, 1961 : 518, fig. u. a-d; 1967 : 600, fig. 28.2. n-p.

HABITAT. Mud at 18 m depth.

RECORD. ML 218 - i.

NOTES. The specimen is an anterior fragment of 25 segments, measuring 7 mm
in length. An achaetous peristomium is followed by 10 thoracic setigers, the first
four of which have capillaries, the remaining six having hooks only. Although
conforming to the description of the South African material, the identity of the
specimen must remain provisional, due to its fragmentary condition.

Notomastus sp.

HABITAT. Sand, 2-9 m depth.

RECORDS. ML 97 - i ; ML 118 - i.

NOTES. Both specimens are anterior fragments consisting of n thoracic setigers
and a few abdominal segments. Setigers i to 4 are triannulate and setigers 5 to n
are quadriannulate. The thoracic segments are strongly areolated, particularly
the more anterior ones. The abdominal tori are well developed. A specific identi-
fication cannot be made with any certainty because of the absence of branchiferous
segments.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 187

Family ARENICOLIDAE
Abarenicola claparedii claparedii (Levinsen, 1883)

Arenicola claparedii: Fauvel, 1927 : 163, fig. 57. k-n.
Abarenicola claparedii: Wells, 1959 : 307, fig. 2.

HABITAT. Coralline silty mud at LWM.
RECORD. Matiu Is. - i.

NOTES. The single specimen was kindly examined by Prof. G. P. Wells who
identified it, from its superficial characters only, as one of the cystless species of
Abarenicola, probably claparedii claparedii. This determination will have to be
confirmed on the basis of its internal anatomy.

DISTRIBUTION. Mediterranean.

Family MALDANIDAE

The following descriptions of the maldanid material are based on notes provided
by Dr. Charlotte Mangum.

Subfamily EUCLYMENINAE
Clymenella (=Macroclymene) sp. 1

HABITAT. Silty shell gravel on Thalassia flat.
RECORD. Graham Pt. - i.

NOTES. The specimen is incomplete (the head and probably five anterior setigers
are missing) and has 48 setigers plus a caudal funnel. The latter carries long and
short cirri, which are arranged in an alternating pattern of one long, two short, one
long, two short and so on, in addition to a median ventral cirrus that is 1-5 to 2-0
times the length of the long cirri. Posterior pre-anal achaetous segments are absent.

Clymenella (=Macroclymene) sp. 2

HABITAT. Under boulder in muddy silt at LWM.
RECORD. Komimbo Bay- i.

NOTES. An anterior fragment composed of 27 setigers is available. It has a
cephalic plaque with nuchal organs extending nine-tenths of its length. Prostomial
pigment spots are absent. There are shallow clefts in the medial and posterior
margins of the cephalic rim. Uncini are present on all setigers including the first
three.

i88 P. E. GIBBS

Clymenella (=Euclymene) sp.

HABITAT. Silty sand with coral debris on lower shore.
RECORD. Komimbo Bay - i.

NOTES. The specimen is complete with 19 setigers plus one or two posterior
achaetous segments. A cephalic plaque is present, the rim of which has six well
defined lobes along the dorsal margin. There are a few prostomial pigment spots.
Setigers i to 3 each have a single large aciculum. A caudal funnel is present and
this carries a median ventral cirrus that is more than 1-5 times the length of the other
cirri. Questionably identified as Praxillella kollikeri (Mclntosh, 1885, as Pr axilla
kollikeri).

Praxillella sp.

HABITAT. Silty fine coral sand at LWM.
RECORD. Matiu Is. - i.

NOTES. The complete specimen has 19 setigers plus two posterior achaetous
segments. A cephalic plaque is developed with nuchal organs extending over nine-
tenths of its length. The cephalic rim has posterior clefts only. Pigment spots on
the prostomium are profuse. There are three or four uncini on setigers i and 2 and
four on setiger 3. A caudal funnel is not present. The anal plug is just protruding
and is surrounded by cirri of equal length. The specimen resembles P. affinis (Sars)
but is probably a new species.

Euclymeninae sp. A

HABITAT. Muddy sand at 9 m depth.
RECORD. ML 56-1.

NOTES. This specimen is an anterior fragment of nine setigers. The cephalic
plaque has nuchal organs extending across three-quarters of its length. Prostomial
pigment spots are absent. The first setiger is not clearly separated from the
prostomium. One pair of acicula is present on setiger i, two pairs on setiger 2 and
one or two pairs on setiger 3, with uncini on the following segments.

Euclymeninae sp. B

HABITAT. Between boulders in muddy silt at MTL.
RECORD. Tetel Is. - i.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 189

NOTES. This incomplete specimen is composed of the prostomium and eight
setigers. The nuchal organs extend for nine-tenths of the length of the cephalic
plaque and there are very shallow clefts around the medial and posterior margins of
the cephalic rim. Pigment spots are present ventrally. Setiger i has one pair of
acicula and setigers 2 and 3 have two pairs, the subsequent setigers having uncini.

Euclymeninae sp. C

HABITAT. Silty sand at 24 m depth.
RECORD. ML 230-1.

NOTES. The specimen is fragmentary, consisting of 15 anterior setigers. The
cephalic plaque is unusual in having pigment spots within the plaque as well as on the
ventral surface. The cephalic rim is very high and has posterior clefts only.

Subfamily NICOMACHINAE
? Nicomache sp.

HABITAT. Coarse coral sand below LWM.
RECORDS. Tetel Is. - 2.

NOTES. The material consists of two large anterior fragments with seven and
eight setigers. They appear to be identical except that one specimen has only large
setae on setigers I to 3 but the other specimen has four small setae on setiger i, a
small (left) and a large (right) seta on setiger 2 and one large seta on either side of
setiger 3. The latter specimen may be abnormal due to aberrant growth or re-
generation. In both specimens the first three setigers are light, the rest being grey-
brown to dark in colour.

Family OWENIIDAE

Myriochele eurystoma Caullery, 1944

(Fig. 15. A)

Myriochele eurystoma Caullery, 1944 : 52, fig. 42. a-d, f.

HABITAT. Silty sand at 35 m depth.
RECORD. ML 72-1.

NOTES. The specimen is an anterior fragment about 15 mm long and has a
diameter of 0-5 mm. It has a large mouth with a prominent buccal organ on its

i go

P. E. GIBBS

ventral margin (fig. 15. A). The prostomium and anterior segments are pigmented
reddish-brown, with two eye-spots situated ventro-laterally. Between the latter,
running over the dorsal surface, there is a thin unpigmented band. The tube is
composed chiefly of sponge spicules cemented with their long axes around the
circumference of the tube, the latter having a diameter of about 1-2 mm.

The specimen is referred to M. eurystoma on account of its large mouth and its
pigmentation. Caullery notes that a specimen from 32 m depth was pigmented but
specimens from deeper waters (131-1570 m) lacked colour. The details of the
posterior region are unknown.

DISTRIBUTION. East Indies.

Myriochele heruensis n. sp.

(Fig. 15. B-G)

DESCRIPTION. Of the fourteen specimens taken at ML 134, only three could
be removed intact from their tubes. The holotype is 7-6 mm long for 18 segments
and the two other complete paratypes measure 6-6 and 7-5 mm for 16 and 17 seg-
ments respectively. The largest specimen probably measured between 10 and 12
mm long when intact. The diameter of the worm is between 0-28 and 0-40 mm.

H G F E

FIG. 15. Myriochele eurystoma. (A) Anterior region. Myriochele heruensis n.sp. (B-D)
Ventral and lateral views of anterior region. (E) Dorsal view of posterior region. (F-G)
Profile and face views of uncinus. Myriochele sp. (H) Ventral view of anterior region.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 191

The prostomium is short with a rounded anterior margin (fig. 15. B). In those
specimens preserved with the prostomium protracted, a shallow groove can be
discerned extending across the dorsal surface of the prostomium, terminating on
either side at the level of ventro-lateral eye-spots (fig. 15. c). In other specimens
the prostomium is contracted, and in these the dorsal groove appears as a distinct
fold (fig. 15. D). This groove or fold is quite marked in all specimens and serves as
the characteristic feature of the species. In addition to the two red eye-spots, two
small patches of red pigment are present on the antero-dorsal surface of the prosto-
mium and there is also a thin red line along the posterior margin of the dorsal groove.

The first three setigers are short. From setiger 4 there is a progressive increase
in the length of the segments, reaching a maximum in the middle body region, and
then successive segments become shorter. The pygidium has two short, stout anal
cirri (fig. 15. E).

The setae are of the usual Myriochele types. Notosetae are slender capillaries
with short lateral processes along either side of the blade. Neurosetae, present from
setiger 4, are uncini, each with two subequal teeth at the apex (fig. 15. F-G). In
the middle body segments, the tori are composed of 7 or 8 vertical rows of uncini.

The tube is composed of shell fragments which are cemented with their flat
surfaces at right angles to the long axis of the tube. The latter has a diameter of
0-8 to i-o mm and a maximum length of about 25 mm.

As noted above the distinct groove across the dorsal surface of the prostomium
is sufficient to distinguish M. heruensis from all other species. This feature may be
interpreted as a feeding adaptation in that it provides a mechanism whereby the
position of the mouth can be changed.

HABITAT. Sand at 16 m depth.

RECORDS. ML 134 - 14.

British Museum (Natural History) Registration No. Holotype 1970-77

Paratypes 1970-78

Myriochele sp.

(Fig. 15. H)

HABITAT. Sand at 5 m depth.
RECORD. ML 229 - i.

NOTES. The specimen lacks posterior segments but probably measured about
12 mm long when intact. It has a diameter of 0-24 mm. The anterior margin of the
prostomium is bluntly rounded and, ventrally, the mouth extends about half the
prostomial length, ending at the level of two red eye-spots which are ventro-lateral
in position (fig. 15. H).

192 P. E. GIBBS

The tube has a diameter of 0-4 mm and consists of sand and shell fragments
cemented with their long axes along the length of the tube.

The specimen shows a general similarity to M. minor Caullery but differs in having
eye-spots.

Owenia fusiformis Delle Chiaje, 1844

Owenia fusiformis : Fauvel, 1927 : 203, fig. 71. a-f; 1953 : 39 1 ' fig- 2O 3- a ~f; Day, 1967 : 649, fig.
31.1. e-j.

HABITAT. Medium and coarse sand, 2-18 m depth.

RECORDS. ML 96-3; ML 118 - 3; ML 134- 5; ML 190 - 2; ML 192 - 13;
ML 203 - i ; ML 204 - 60 ; ML 229 - 2 ; ML 283 - i.

DISTRIBUTION. Cosmopolitan.

Family SABELLARIIDAE
Lygdamis ehlersi (Caullery, 1913)

Tetreres ehlersi Caullery, 1913 : 201, figs a-d.
Lygdamis ehlersi: Caullery, 1944 : 62, figs 49-50.

HABITAT. Silty sand at LWM.
RECORDS. Fintry Pt. - 5.

NOTES. The largest specimen is 50 mm long for 50 segments. The tube is
constructed of gravel, shell and calcareous alga fragments. The chief characters
include the following: on the operculum, there are 19 or 20 external and 6 or 7
internal paleae on each side and 10 groups of buccal cirri. The median cirrus (lobe
Preoral) is stout, longer than the palps, and has a truncated tip which is darkly
pigmented a bluish-black colour. There are 13 to 15 pairs of branchiae.

Comparing these specimens with the type description they have fewer external
(20 compared to 25) and internal (6 or 7 instead of 12) paleae but correspond in all
other details.

Although Hartman (1959) indicates that L. ehlersi may be a synonym of L. indicus
the two species appear to be distinct and separable on the form of the median cirrus.
In L. ehlersi the median cirrus is stout, longer than the palps and has a truncated
tip while in L. indicus this organ is shorter than the palps and tapers to a point.

DISTRIBUTION. East Indies.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 193

Lygdamis indicus Kinberg, 1867

Lygdamis indicus: Johansson, 1925 : 8, fig. 2. 2-7; Fauvel, 1953 : 399, fig. 209. a-k; Day, 1967 :
677. fig- 33-3- c-h.

HABITAT. Undersurface of coral boulder at LWM.
RECORD. Pirikale Is. - i.

NOTES. The specimen is 30 mm long (diameter = 4 mm) with 48 setigers. It
has 37 external and 17 internal paleae on each side. The tube is a fragile structure
constructed of shell fragments.

DISTRIBUTION. Tropical Indo-west-Pacific.

Family PECTINARIIDAE
Pectinaria (Pectinaria) antipoda Schmarda, 1861

Pectinaria (Pectinaria) antipoda Schmarda, 1861 : 46, pi. 24, fig. 199, with text-figs; Pruvot,
1930 : 78, pi. 3, figs 93-95; Fauvel, 1953 : 403, fig. 211. e-g.

HABITAT. Mud and sand, 4-22 m depth.
RECORDS. ML 37 - i ; ML 203 - i ; ML 229 - i.

NOTES. All three specimens are too small (less than 10 mm) to be certain of their
identity but they appear to belong to this Indo-west-Pacific species.

DISTRIBUTION. Persian Gulf; West Pacific.

Family AMPHARETIDAE

Subfamily MELINNINAE
Isolda pulchella Miiller, 1858

Isolda sibogae Caullery, 1944 : 102, fig. 83. a-h.

Isolda pulchella: Day, 1963 : 434 (synonymy); 1967 : 691, fig. 35.1. k-n.

HABITAT. Mud and silty sand, 2-18 m depth.
RECORDS. ML 56- i; ML 196- i; ML 218- i.
DISTRIBUTION. Atlantic and Indian Oceans; East Indies.

194

P. E. GIBBS

Subfamily AMPHARETINAE
? Sosane wireni Caullery, 1944

(Fig. 16. A-B)

Sosane wireni Caullery, 1944 : 87, fig. 69. a-e.
HABITAT. Silty sand at 24 m depth.
RECORD. ML 230 - i.

NOTES. The specimen is 7 mm long for 26 setigers of which 15 are thoracic and
ii abdominal. It is referred to S. wireni on account of the branchiae being arranged
with three pairs in a transverse line across setiger i and a smaller fourth pair on
setiger 2. However there is some doubt as to the identity since Caullery omits any
reference to the diagnostic features of the genus, namely, the presence of (i) a
nephridial papilla on the branchial ridge and (ii) specialised notosetae on elevated
posterior notopodia (Day, 1964). There does not appear to be a nephridial papilla
present on the Solomon Islands specimen but the notopodia of setiger 13 are elevated
(fig. 16. A) and each carries about 20 modified notosetae which have a 'penicillate'
appearance (fig. 16. B).

DISTRIBUTION. East Indies.

FIG. 16. ? Sosane wireni. (A) Elevated notopodium of setiger 13.
(B) Modified notosetae from same setiger.

Family TEREBELLIDAE

Subfamily TRICHOBRANCHINAE

Terebellides stroemi Sars, 1835

Terebellides stroemi: Fauvel, 1927 : 291, fig. 100. i-q; 1953 : 436, fig. 231. i-q; Day, 1967 : 713,
fig. 36.1. f-j.

HABITAT. Mud and silty sand, 2-24 m depth.

RECORDS. ML 155 - 31 ; ML 157 - 3 ; ML 196 - i ; ML 218 - i ; ML 230 - 3.

DISTRIBUTION. Cosmopolitan.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 195

Subfamily POLYGIRRINAE
Amaeana trilobata (Sars, 1863)

Amaea trilobata: Fauvel, 1927 : 285, fig. 99. a-e.
Amaeana trilobata: Day 1967 : 718, fig. 36.3. e-h.

HABITAT. Mud, 11-22 m depth.

RECORDS. ML 29 - i ; ML 194 - i.

DISTRIBUTION. North Atlantic; Mediterranean; southern Africa; Japan.

Lysilla ubianensis Caullery, 1944

Lysilla ubianensis Caullery, 1944 : 197, fig. 156. a-e; Day, 1967 : 721, fig. 36.3. i-j.

HABITAT. Silty mud, MTL to LWM, and at 13 m depth
RECORDS. Tetel Is. - 3 ; Komimbo Bay - i ; ML 69-1.
DISTRIBUTION. East Africa; East Indies.

Subfamily THELEPINAE
Euthelepus kinsemboensis Augener, 1918

Euthelepus kinsemboensis Augener, 1918 : 548, pi. 6, fig. 161, pi. 7, fig. 250, text-fig. 95; Fauvel,
I 93 0: 553. fig- 9- a-f; 194? : 7 8 - fi g- 75- a'-d'; Day, 1967 : 726, fig. 36.5. e-i.

HABITAT. Under coral boulder on reef platform.
RECORD. Graham Pt. - i.

NOTES. The specimen is an anterior fragment of 34 setigers which corresponds
in detail to the specimen from New Caledonia described by Fauvel except that the
branchial filaments, situated on segments 2, 3 and 4, number 10, 8 and 6 respectively,
not 12, 8 and 4. The type specimen from Angola has only 12 branchial filaments
instead of 24, but according to Fauvel (1930) this variation is attributable to the
difference in the size of the specimens.

DISTRIBUTION. Angola; New Caledonia.

Subfamily TEREBELLINAE
Eupolymnia nebulosa (Montagu, 1818)

Polymnia nebulosa: Fauvel, 1927 : 257, fig. 89. a-g; 1953 : 419, fig. 219. a-g.
Eupolymnia nebulosa: Imajima & Hartman, 1964 : 337; Day, 1967 : 744, fig. 36.9. f-h.

HABITAT. Under coral boulder on silty sand at LWM.

RECORD. Graham Pt. - i.

DISTRIBUTION. Atlantic; South Africa; Mediterranean; Indo-west Pacific.

ig6 P. E. GIBBS

Loimia medusa (Savigny, 1820)
Loimia medusa: Fauvel, 1953 : 4 J 6, fig- 218. a-f; Day, 1967 : 743, fig. 36.9. a-e,

HABITAT. A wide variety of deposits, silty mud to Acropora rubble, from above
MTL to LWM on reef platforms; in crevices within reef platforms and amongst
encrustations on wharf piles ; silty sand, 9-24 m depth.

RECORDS. Tetel Is. - 3 ; Komimbo Bay - 9 ; Mamara Pt. - i ; Matiu Is. - i ;
Lauvie Is. - 6 ; Fintry Pt. - 9 ; Yandina - 2 ; Maramasike Pg. - i ; ML 69 - i ;
ML 96 - i ; ML 230 - i.

NOTES. The largest specimens, up to 40 cm in length with over 200 segments,
were found in coral debris deposits (chiefly Acropora fragments) on reef platforms.
Their tubes were loosely constructed of Halimeda, coral and shell fragments and
extended to a depth of over 30 cm. Although conspicuous on account of their
white feeding tentacles spread over the surface, specimens are difficult to capture
in this habitat because of their ability to rapidly withdraw to the bottom of their
deep tubes when disturbed. However smaller specimens which are commonly
found under coral boulders lying in silty mud are easily taken.

DISTRIBUTION. Cosmopolitan in temperate and tropical waters.

Pista dibranchis n. sp.

(Fig. 17. A-E)

DESCRIPTION. A specimen from Tetel Island has been selected for the holotype.
It is incomplete and measures 12 mm for 32 segments. The largest of the paratypes
is 40 mm long for 130 segments. The body diameter is about i-o mm.

The tentacular lobe bears numerous tentacles and is without eye-spots. Semi-
circular lateral lobes are present on segments 2 and 3. A pair of stalked branchiae
arises from the anterior margin of the second segment and each has numerous
filaments arranged spirally, with a maximum of 6 or 7 whorls, around the central
stalk (fig. 17. A) ; often one of the pair is much larger than the other. In larger
specimens the nephridial papillae on segments 6 and 7 (setigers 3 and 4) are cons-
picuous.

Notosetae commence on segment 4 and extend to segment 20 (setiger 17). Each
bundle consists of broad-winged, smooth-tipped capillaries of two types, namely, a
longer stout-shafted form (fig. 17. B) and a shorter, slender form with a recurved tip
(fig. 17. c). There are usually about n of each type in a bundle. Uncini start on
segment 5 (setiger 2) and are arranged in single rows on the anterior segments,
becoming alternate in the posterior thorax. Uncini are of similar form throughout
the thorax and each has 5 to 7 teeth surmounted by 15 to 20 smaller teeth above the
main fang (fig. 17. D, E). The dental formula is thus: MF : 5-7 : 15-20. The uncini
lack basal prolongations or shafts, even in the first row. Abdominal uncini are
borne on short, square pinnules.

The tube is composed of sand grains cemented together with mud particles.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS

197

At first it was assumed that these specimens possessing only one pair of branchiae
were P. typha juveniles in which the second pair of branchiae was either undeveloped
or had been lost. However subsequent examination revealed that the uncini of the
anterior segments lacked the basal shafts found in P. typha, and also some of the
specimens contain coelomic oocytes, apparently approaching a mature condition.

In having only one pair of branchiae, P. dibranchis resembles P. vinogradovi
Uschakov but lacks the transverse dorsal membrane between the branchial stalks
that is present in the latter species. According to Uschakov (1955), P. cristata
(Miiller) may also have one pair of branchiae but this species differs in possessing
shafted uncini. In terms of the structure of the branchial apparatus, P. dibranchis
is intermediate between P. unibranchia Day, which has a single branchia, and P.
typha Grube, which has two pairs, all three species having branchiae with spirally
arranged, or whorled, filaments.

HABITAT. A wide variety of deposits, silty mud to coarse sand and shell gravel,
from MTL to 24 m depth.

RECORDS. Haroro - i ; Tetel Is. - 4 (including holotype) ; Graham Pt. - i ;
Fintry Pt. - 3 ; ML 38 - i ; ML 204 - i ; ML 230 - i ; ML 283 - 6; ML 296 - i.

British Museum (Natural History) Registration No. Holotype 197079

Paratypes 1970-80-89

NOTE. The habitat of P. dibranchis overlaps that of P. typha ; on Tetel Island the
two species were taken from the same silty mud deposit on the reef platform at about
MTL.

FIG. 17. Pista dibranchis n.sp. (A) Dorsal view of anterior region.
(B-C) Thoracic notosetae. (D-E) Profile and face views of uncinus.

ig8 P. E. GIBBS

Pista typha (Grube, 1878)

Terebella (Pista) typha Grube, 1878 : 232, pi. 12, fig. 4.
Pista typha: Fauvel, 1953 : 424, fig. 222. a-c.

HABITAT. Silty sand and mud, MTL to LWM.
RECORDS. Tetel Is. - 23; Fintry Pt. - i.
DISTRIBUTION. Indo-west-Pacific.

Reteterebella queenslandia Hartman, 1963

Reteterebella queenslandia Hartman, 1963 : 355, figs 1-3.

HABITAT. Acropora rubble at LWM.
RECORDS. Tetel Is. - i ; Maraunibina Is. - i.

NOTES. The genus Reteterebella Hartman differs from the closely allied genus
Eupolymnia Verrill in lacking notosetae on segment 4, thus having 16 instead of 17
thoracic setigers, and also in the first appearance of the double rows of uncini on the
sixth (segment 10 or setiger 6), not the seventh uncinigerous segment (= setiger 8 in
Eupolymnia).

DISTRIBUTION. North-east Australia (Queensland).

Terebella ehrenbergi Grube, 1870

Terebella ehrenbergi: Fauvel, 1953 : 421, fig. 220. a-c; Day, 1967 : 748, fig. 36.10. g-i.

HABITAT. Crevices in reef platform.

RECORDS. Matiu Is. - 2.

DISTRIBUTION. Tropical Indo-west-Pacific. .

Family SABELLIDAE
Branchiomma cingulata (Grube, 1870)

Dasychone cingulata: Willey, 1905 : 308, pi. 7, figs 170-173; Fauvel, 1953 : 442, fig. 234. f-h.
Branchiomma cingulata: Johansson, 1927 : 161, text-fig. 14.1-2; Imajima & Hartman, 1964 : 355.

HABITAT. Embedded in Acropora at LWM.
RECORD. Tetel Is. - i.

NOTES. The specimen is small, about 7 mm long. It has six radioles on each
side of the branchial crown and the thoracic region consists of five segments.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 199

Specimens from Ceylon and Japan have a greater number of radioles and eight
thoracic segments but are larger-sized, which may account for these differences.
DISTRIBUTION. Indo-Pacific.

Hypsicomus phaeotaenia (Schmarda, 1861)

Hypsicomus phaeotaenia : Fauvel, 1953 : 447, fig. 236. a-1; Day, 1967 : 761, fig. 37.2. i-n.
HABITAT. Embedded (boring?) in coral, especially Forties, and in beachrock.

RECORDS. Kokomtambu Is. - 1 ; Tetel Is. - 4 ; Komimbo Bay - 3 ; Mamara Pt. - 1 ;
Matiu Is. - 5 ; Paleki Is. - i ; Lauvie Is. - 7; Graham Pt. - 3.

DISTRIBUTION. Western Africa; Mediterranean; Indo-west-Pacific.

Megalomma intermedium (Beddard, i

Branchiomma intermedium Beddard, 1888 : 261, pi. 21, figs 4-7; Fauvel, 1953 : 444, fig. 234. e.
Megalomma intermedium: Pillai, 1965 : 164.

HABITAT. Embedded in Porites boulder towards LWM.

RECORD. Tetel Is. - i.

DISTRIBUTION. Mergui Archipelago; Philippine Islands.

Megalomma linaresi (Rioja, 1918)

Branchiomma linaresi: Fauvel, 1927 : 317, fig. no. a-1.

HABITAT. Embedded in Acropora at LWM and at 5 m depth.

RECORDS. Tetel Is. - i ; Yandina - i.

NOTES. The larger specimen is 45 mm long, including the branchial crown which
has a length of 10 mm. There are 16 to 19 radioles on each side. The dorsal radioles
carry the largest subterminal eyes, which gradually diminish in size on the lateral
radioles and are absent from the ventral ones. The collar gapes widely on the
dorsal side: it has lateral notches and large ventral lobes. There are 8 thoracic
segments, the notosetae of which consist of narrow-winged capillaries and sub-
spatulate setae.

M. linaresi is known from northern Spain and the Mediterranean. Despite the
absence of records from the Indian Ocean, the specimens from the Solomon Islands
appear to be identical.

DISTRIBUTION. North Atlantic (Spain) ; Mediterranean.

200 P. E. GIBBS

Megalomma quadrioculatum (Willey, 1905)

Branchiomma quadrioculatum Willey, 1905 : 307, pi. 7, figs 168-169.
Megalomma quadrioculatum: Day, 1967 : 758, fig. 37.1. h-o.

HABITAT. Amongst Phyllochaetopterus socialis tubes on reef platform towards
LWM ; amongst serpulid tubes on undersurface of coral boulder.

RECORDS. Cape Esperance - 36; Komimbo Bay - i.

NOTES. The specimens were removed from sandy tubes constructed amongst
the sponges and other encrusting organisms around the bases of the chaetopterid
tubes. The larger specimens measure 13 to 14 mm long. The thorax consists of
6 or 7 segments with notosetae of two types, namely narrow-winged capillaries and
paleae. There are 6 to 9 radioles on each side of the branchial crown and, in most
specimens, only the two dorsal radioles carry subterminal eyes. However, a few
specimens have smaller eyes on one or two radioles besides the dorsal radioles.

The type specimen from Ceylon (23-5 mm in length) is almost twice the size of the
Solomon Islands specimens and has 14 radioles per branchial lobe with subterminal
eyes on the four dorsal radioles. It would seem therefore that the second (and
occasionally a third) pair of eyes develops at a later stage and, until acquired, M.
quadrioculatum bears a superficial resemblance to M . bioculatum (Ehlers) from which
it can be distinguished by the presence of paleae in the notopodia.

DISTRIBUTION. Indian Ocean.

Megalomma trioculatum Reish, 1968

Megalomma trioculatum Reish, 1968 : 226, fig. 5. i-io.

HABITAT. Sand, 2-9 m depth.
RECORDS. ML 96 - i; ML 296- i.

NOTES. The two specimens measure 15 to 16 mm long, the branchial crown
being 2-0 to 2-5 mm in length. Both have 6 thoracic setigers (instead of 8 or 9) and
the collar gapes widely on the dorsal side of the first setiger instead of extending to
the mid-segmental line. Otherwise they agree closely with type description.

At the bases of the two dorsal radioles, there are longitudinal streaks of an orange-
brown pigment and the radioles are banded a similar colour. The subterminal eyes
are brown or mauve. The tube is a fragile structure coated with sand grains.

DISTRIBUTION. Marshall Islands.

Megalomma vesiculosum (Montagu, 1815)

Branchiomma vesiculosum: Fauvel, 1927 : 315, fig. 109. a-q.
Megalomma vesiculosum: Day, 1967 : 758, fig. 37.1. p-u.

HABITAT. Muddy sand between MTL and LWM.
RECORDS. Tetel Is. - 20.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 201

NOTES. The subterminal eyes are entire, not divided as in M. pacificum Johansson
although the latter may be a synonym of M. vesiculosum, according to Mesnil &
Fauvel (1939).

DISTRIBUTION. Atlantic Ocean; Mediterranean; Indian Ocean.

Potamilla ehlersi Gravier, 1906

Potamilla ehlersi Gravier, igo6a : 37; 1908 : 87, pi. 6, fig. 260-264; Fauvel, 1953 : 449,
fig. 238. g-i.

HABITAT. Amongst Phyllochaetopterus socialis tubes on reef platform towards
LWM.

RECORD. Cape Esperance- i.

NOTES. The specimen is 13 mm long (branchial crown = 3 mm). There are
nine radioles on each branchial lobe ; the dorsal ones are without eyes but these are
present on the next 4 or 5 radioles. There are n segments in the thorax, the
anterior half of which is coloured chocolate-brown on the dorsal side. The tube is
membraneous, covered with sand grains, and, before dissection, its aperture was
rolled, as found in P. reniformis (Muller) (cf. Fauvel, 1927, fig. 107 1).

DISTRIBUTION. Tropical Indo-west-Pacific.

Sabellafusca Grube, 1870

Sabella fusca: Fauvel, 1927 : 302, fig. 104. a-f; Day, 1967 : 764, fig. 37.2. t-v.
HABITAT. Beachrock.
RECORDS. Lauvie Is. - 3.
DISTRIBUTION. Mediterranean; tropical Indo-west-Pacific.

Sabella melanostigma Schmarda, 1861

Sabella melanostigma Schmarda, 1861 : 36, pi. 22, fig. 190, with text-figs; Fauvel, 1953 : 439,
fig. 232. h-n.

HABITAT. Mud and silty sand, 24-33 m depth.
RECORDS. ML 100 - i ; ML 230 - 5.

NOTES. The specimens are only about 12 mm long and were extracted from
narrow tubes (0-65 mm diameter) composed of mud particles. They are referred to

P. E. GIBBS

this species because they possess the peculiar 'shovel-like' pick-axe setae. However
pigment spots above the parapodia are lacking. There are 7 radioles, each with 2
or 3 pairs of eye-spots, on either branchial lobe. The ventral lobes of the collar are
large and there is a wide gap between the dorsal lobes.

DISTRIBUTION. Circumtropical.

Sabellastarte sanctijosephi (Gravier, 1906)

Eurato sancti-josephi Gravier, igo6a : 42; 1908 : 105, pi. 7, figs 281-283, pi. 8, figs 284-285.
Sabellastarte indica: Fauvel, 1953 : 445, fig. 235. a-h.
Sabellastarte sanctijosephi: Day, 1967 : 771, fig. 37.5. f-i.

HABITAT. In crevices in Forties boulders at LWM (Gibbs, 1969, fig. 137).

RECORDS. Tetel Is. - i ; Paleki Is. - 5.

DISTRIBUTION. Western Africa; tropical Indo-west-Pacific.

Family SERPULIDAE

Subfamily SERPULINAE

Hydroides minax (Grube, 1878)

Serpula minax Grube, 1878 : 269, pi. 15, fig. 5.

Hydroides minax: Fauvel, 1953 : 4^o, fig. 241. f; Pillai, 1960 : 8, text-fig. 3. a-e.

HABITAT. Amongst Phyllbchaetopterus socialis tubes at LWM.
RECORDS. Matiu Is. - 2.

NOTES. These specimens were referred to H. monoceros Gravier, a closely allied
species, in an earlier paper (Gibbs, 1969).
DISTRIBUTION. Ceylon; Philippine Islands.

Hydroides uncinata (Philippi, 1844)

Hydroides uncinata: Fauvel, 1927 : 357, fig. 122. a-h; Day, 1967 : 805, fig. 38.4. h-i.
HABITAT. On Acropora at 5 m depth.
RECORD. Yandina-i.
DISTRIBUTION. Atlantic Ocean; Mediterranean; East Africa; Japan.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 203

Mercierella enigmatica Fauvel, 1923

Mercierella enigmatica: Fauvel, 1927 : 360, fig. 123. a-o; Day, 1967 : 812, fig. 38.5. o-s.
Neopomatus uschakovi Pillai, 1960 : 28, pi. i, figs 1-2; text-figs 10. h, n. a-h, 12. a-h; Straughan,
1966 : 139 (synonymy).

HABITAT. Brackish water - on stones and in sponge.

RECORDS. Komimbo Bay (freshwater creek) - num. ; mouth of Lunga River at
Lunga Pt. - 28.

NOTES. In a study of the brackish water serpulids along the east coast of
Australia, Straughan (1966) discovered that there is a continuous cline in the characters
used to separate the two genera Mercierella Fauvel and Neopomatus Pillai. The
Mercierella form is characteristic of the southern populations around Sydney and
the warm water Neopomatus form is typical for the Queensland populations.

The material from Guadalcanal is typical Neopomatus in that there are one or
two rows of outwardly directed spines on the operculum, the collar and thoracic
membranes are fused dorsally for up to three-quarters of the length of the thorax,
and the collar setae have modified proximal teeth.

DISTRIBUTION. Cosmopolitan in estuarine water.

Serpula hartmanae Reish, 1968

Serpula hartmanae Reish, 1968 : 228, fig. 5. 11-16.

HABITAT. Attached to coral on reef platform.
RECORDS. Mamara Pt. - i ; Komimbo Bay - 3.
DISTRIBUTION. Marshall Islands.

Serpula vermicularis Linnaeus, 1767

Serpula vermicularis: Fauvel, 1927 : 351, fig. 120. a-q; 1953 : 454, fig. 239. a-q; Dew, 1959 : 22,
fig. 3. a-h; Day, 1967 : 809, fig. 38.5. a-h.

HABITAT. Attached to coral boulders on reef platform.
RECORDS. Tetel Is. - 2 ; Komimbo Bay - 5.
DISTRIBUTION. Cosmopolitan.

Spirobranchus giganteus (Pallas, 1766)

Spirobranchus giganteus : Fauvel, 1953 : 462, fig. 242. a-g; Day, 1967 : 803, fig. 38.3. h-k.

HABITAT. Embedded in living Porites boulders (Gibbs, 1969, fig. 138) and
encrusting on reef platform.

204 P. E. GIBBS

RECORDS. Kokomtambu Is. -2; Tetel Is. - v. num.; Komimbo Bay -num.;
Mamara Pt. - i ; Matiu Is. - 9; Paleki Is. - v. num. ; Sifola - i ; Graham Pt. - num.

NOTES. This species is common in moderately sheltered conditions and the
largest specimens, over no mm in length and 10 mm in diameter, were extracted
from living Porites at Paleki Island. In comparison, encrusting forms on dead
coral are much smaller in size.

DISTRIBUTION. West Indies; Indo-west-Pacific.

Spirobranchus coutierei (Gravier, 1908)

Pomatoceropsis coutierei Gravier, 1908 : 125, pi. 8, figs 294-299, text-figs 482-487.

HABITAT. Embedded in Porites towards LWM.
RECORDS. Tetel Is. - 3.

NOTES. Following Straughan's (1967) key to the Indo-west-Pacific species, this
identification is based on the following two characters - (i) the operculum has three
processes which are deeply divided so as to give six distinct horns, and (ii) the
interbranchial membrane is fimbriated.

DISTRIBUTION. East Africa; northern Australia (Straughan, 1967^).
Vermiliopsis glandigerus Gravier, 1908

Vermiliopsis glandigerus Gravier, 1908 : 121, pi. 8, figs 290-291, text-figs 476-481; Fauvel,
1953 : 467, fig. 242. k; Day, 1967 : 813, fig. 38.6. g-i.

HABITAT. Encrusting on reef platform.

RECORD. Matiu Is. - i.

DISTRIBUTION. Western and southern Africa; Indo-west-Pacific.

Subfamily FILOGRANINAE
Filograna implexa Berkeley, 1835

Salmacina dysteri: Fauvel, 1927 : 377, fig. 129. c-k; 1953 : 476, fig. 250. c-k; Dew, 1959 : 50,

fig. 19. a-g.
Filograna implexa: Fauvel, 1927 : 376, fig. 129. a-b; Day, 1955 : 450 (synonymy); 1967 : 817,

fig. 38.7. a-h.

HABITAT. Encrusting on an oyster shell attached to wharf pile.
RECORDS. Yandina - num.

NOTE. No individual with an operculum developed at the end of a branchial
filament was noted amongst the colony; the specimens thus correspond to the
Salmacina form.

DISTRIBUTION. Cosmopolitan.

THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 205

ADDENDUM

Recently Professor J. E. Morton forwarded a small collection of polychaetes which
he collected in the Solomon Islands during the 1965 Expedition. This material
includes specimens of a further two species, bringing the total number of recorded
species to 222.

The species are the following :

Family NEREID AE
Perinereis neocaledonica Pruvot, 1930

Perinereis neocaledonica Pruvot, 1930 : 50, pi. 3, figs 77-79, text-fig. 4. a-c; Fauvel, 1932 : 107;
1953 : 2ii, fig. 108. c-g; Tampi & Rangarajan, 1964 : 106.

RECORDS. Batuona Is - 10.
DISTRIBUTION. Tropical Indo- west-Pacific.

Family GLYCERIDAE
Glycera subaenea Grube, 1878

Glycera subaenea Grube, 1878 : 184, pi. 8, fig. 8; Fauvel, igiga : 425, pi. 16, figs 48-51; Hartman
& Imajima, 1964 : 164; Day, 1967 : 363, fig. 16.3. k-n.

RECORDS. Komimbo Bay - 3.
DISTRIBUTION. Indo-west-Pacific.

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210 P. E. GIBBS

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THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 211

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Dr. P. E. GIBBS

MARINE BIOLOGICAL ASSOCIATION OF THE U.K.

CITADEL HILL

PLYMOUTH

A LIST OF SUPPLEMENTS
TO THE ZOOLOGICAL SERIES

OF THE BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

1. KAY, E. ALISON. Marine Molluscs in the Cuming Collection British Museum
(Natural History) described by William Harper Pease. Pp. 96; 14 Plates.
1965. (Out of Print.) 3.75.

2. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier and
Valenciennes. Pp. 180; u Plates, 15 Text-figures. 1967. 4.

3. TAYLOR, J. D., KENNEDY, W. J. & HALL, A. The Shell Structure of Mineralogy
at the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125; 29 Plates,
77 Text-figures. 1969. 4.50.

4. HAYNES, J. R. Cardigan Bay recent Foraminifera (Cruises of the R.V. Antur)
1962-1964. (In press.)

Printed in England by Staples Printers Limited at their Kettering, Northants, establishment

LIZARDS AND SNAKES FROM

TRANSJORDAN, RECENTLY

ACQUIRED BY THE BRITISH

MUSEUM (NATURAL HISTORY)

Y. L. WERNER

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 21 No. 6

LONDON: 1971

LIZARDS AND SNAKES FROM TRANSJORDAN,

RECENTLY ACQUIRED BY THE
BRITISH MUSEUM (NATURAL HISTORY)

YEHUDAH LEOPOLD WERNER

The Hebrew University of Jerusalem

Pp. 213-256; 6 Plates, 8 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 21 No. 6

LONDON: 1971

THE BULLETIN OF THE BRITISH MUSEUM

(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical series.

Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.

In 1965 a separate supplementary series of longer
papers was instituted, numbered serially for each
Department.

This paper is Vol. 21, No. 6 of the Zoological series.
The abbreviated titles of periodicals cited follow those
of the World List of Scientific Periodicals.

World List abbreviation
Bull. Br. Mus. nat. Hist. (Zool.)

Trustees of the British Museum (Natural History), 1971

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued 3 December, 1971 Price 2

LIZARDS AND SNAKES FROM TRANSJORDAN,

RECENTLY ACQUIRED BY THE
BRITISH MUSEUM (NATURAL HISTORY)

By YEHUDAH L. WERNER

CONTENTS

Page

SYNOPSIS ........... 215

INTRODUCTION. .......... 215

LOCALITIES ........... 216

METHODS ........... 218

LACERTILIA ........... 219

Gekkonidae .......... 219

Agamidae .......... 222

Chamaeleonidae ......... 232

Lacertidae .......... 234

Scincidae .......... 241

OPHIDIA ............ 243

Colubridae . . . . . . . . . . 243

DISCUSSION ........... 246

General biogeography of Transjordan ..... 246

Distribution of reptiles ........ 249

ACKNOWLEDGEMENTS . . . . . . . . .251

APPENDIX ........... 251

REFERENCES ........... 254

SYNOPSIS

A report on 45 lizards and snakes, representing 23 species and subspecies, collected in northern
and southwestern Transjordan, mostly during 1963-1965. Taxonomic characters are pre-
sented, and compared with data from adjacent areas, mainly Cisjordan. Relevant Trans-
jordanian specimens in the Hebrew University of Jerusalem are also considered, and some
identifications are revised. Field observations are cited. Agama pallida haasi ssp. nov. is
described (type: BM 1965.800; 18 paratypes in BM, HUJ, FMNH). The only additions, on
the species level, to the Trans Jordanian fauna, are Coluber rhodorhachis Jan and Malpolon
moilensis Reuss. The ecological and phytogeographical subdivision of Transjordan into
Mediterranean, Irano-Turanian, and Saharo-Sindian territories is reviewed. The distribution of
reptiles appears to accord with this subdivision. The difference between the herpetofaunas of
Trans- and Cisjordan, on the specific and subspecific levels, is greater in the south than in the
north. Notably 7 Irano-Turanian and Saharo-Sindian forms of Transjordan do not occur in
Cisjordan. It is suggested that the Wadi 'Arava together with the steep mountains bordering
it on the east, may constitute a barrier to the distribution of reptiles.

INTRODUCTION

TRANSJORDAN, or Eastern Palestine, is of great zoo-geographical interest. In the
north-west it borders on the mesic (Mediterranean) regions of Cisjordan (or Western
Palestine), and Syria. To the north and north-east its steppe is continuous with the

216 Y. L. WERNER

steppes of Syria and Iraq, while its south-eastern portions are part of the Arabian
desert. In the south-west, along the Wadi 'Arava, Transjordan adjoins the arid
south of Cisjordan (Negev of Isreal), which, through Sinai, affords communication
with north-eastern Africa.

Despite the efforts of numerous naturalists, zoologists and herpetologists, the
herpetofauna of this whole region remains imperfectly known. The best-known
territory is the part of Cisjordan which has been within Israel since 1948, although
the latest review of its herpetofauna in a European language (Haas, 1951) is now
outdated due to subsequent collecting. More recent information is available to
readers of Hebrew (Barash and Hoofien, 1956; Wahrman, 1963; Y. L. Werner, 1966).
The herpetofauna of Sinai was reviewed by Schmidt and Marx (1956) and Marx
(1968), and that of Iraq by Khalaf (1959). The herpetofauna of Syria and Lebanon
was the subject of several older reports (referred to by Flower, 1933; Schmidt, 1939;
Haas, 1951), and one recent publication (Zinner, 1967).

The least known territory, herpetologically, is Transjordan, where little collecting
has been done. Apparently the only recent papers dealing specifically with the
herpetofauna of this area are those of Schmidt (1930), Parker (1935), Haas (1943,
1951), Hoofien (1965, 1969) and Werner (1968). A few reports of broader scope also
deal with Transjordanian reptiles; notably those of Peracca (1894), Barbour (1914),
Schmidt (1939) and Wettstein (1951), and those cited by these authors or by Parker
(1935) and Haas (1943, 1951).

Recently the British Museum (Natural History) obtained 39 specimens of lizards,
and 6 of snakes, from Transjordan, thanks to the thoughtfulness of three parties
whose primary object had not been the collection of preserved reptiles: Mr. S.
Bisserot of the British Jordan Expeditions 1963 and 1965 (Mountfort, 1965), Mr. D.
Western of the University of Leicester, and (one specimen) Mr. W. Larmuth. The
present material makes a notable addition to our knowledge of the reptiles of Trans-
Jordan and their distribution.

In this paper all but one of .these new specimens* are described and discussed
with a view to stimulating the interest of herpetologists in this little-known region.
Comparisons are made with specimens previously collected in Transjordan by Prof.
G. Haas, Prof. H. Mendelssohn and Mr. J. H. Hoofien (Haas, 1943, 1951 ; Hoofien,
1957) and deposited at the Hebrew University of Jerusalem, and with series from
Cisjordan in the same collection. The zoogeographical implications of the limited
data available are discussed.

LOCALITIES

The localities are indicated by numbers in text-figures i and 2, as follows :

1. Jordanian-Syrian border, 5. Wadi Ratam.
Jerusalem-Damascus road. 6. El Azraq.

2. Tell el Mukheizin. 7. Wadi Aseikhim.

3. Ain el Enoquiya. 8. Azraq Shishan.

4. Azraq Druz. 9. Jebel Uweinid.

*A specimen of Stenodactylus grandiceps Haas, $, BM 1963.665, collected by S. Bisserot at the Azraq
Oasis (in a sandy area), was received by the British Museum but not examined by the author.

LIZARDS AND SNAKES FROM TRANSJORDAN

217

10. Qasr Amra.

11. Tell Qarma.

12. Qa el Umari.

13. Shaubak.

14. Wadi Musa.

15. Petra.

16. Basta.

17. Rum.

18. Aqaba.

-33

-32

-31

-30;

\ _
\
\
\

\
\

AMMANo

12.

FIG. i. Localities from which material is reported here. The rectangle around 6 is en-
larged in fig. 2. Locality names in text. (Political frontiers as before 1966.)

218

Y. L. WERNER

FIG. 2. Localities near Azraq from which material is reported here. This area corresponds
to the rectangle around 6 in fig. i. Locality names in text. B, Basalt areas; W, Water;
MF, Mud flats.

METHODS

Abbreviations:

BM British Museum (Natural History), London.

FMNH Field Museum of Natural History, Chicago.

HUJ-R Herpetological Collection, The Hebrew University of Jerusalem.

SMF Senckenberg Museum, Frankfurt.

SV Snout-vent length.

%SV Percentage of snout-vent length.

Measurements: SV length is from the tip of the snout to the anterior margin of
the cloaca. Measurements of parts of the body are preferably expressed in %SV
(see abbreviations, also Werner, 1969) rather than in absolute values. In Agama
pallida, head length to occiput is from the tip of the snout to the palpated occipito-
atlantal articulation, in a straight line (with dividers); and head length, total is
from the tip of the snout to the rear of the angle of the jaws, parallel to the long
axis of the animal. The head index is derived from the formula 100 x "Head
length, total"/"Head width".

In the case of Agama pallida, the statistical significance of differences in numerical

LIZARDS AND SNAKES FROM TRANSJORDAN 219

values between sample pairs was tested in the following manner: i. The signific-
ance (a = 0-05) of the difference between the variances of the two samples was
checked, using the F-distribution. 2. Regardless of the result, the statistic t
(Winer, 1962 : 28) was computed, and used in Student's -test (Winer, 1962 : 28-29)
as follows: 3. First, t was assigned n a -f n b 2 degrees of freedom; all cases in
which sample means did not differ significantly, needed no further consideration.
4. Where sample means differed significantly (a = 0-05), and the variance had not
differed significantly, the result was accepted. 5. Where the means differed
significantly, but the variances had also differed significantly, the significance of
the difference between the means was confirmed by assigning to t only the degrees
of freedom of the smaller of the two samples, making use of Welch's approximation
(Winer, 1962 : 37-3$) to the i' distribution, (X a - X b ) - (H a - ft,)-

V(s'/n a ) + (s2/n b )
Field, notes are credited expressly to the observer (collector) except where the

observed animal is cited by its number, whereby the collector's name can be located

in 'Material Examined'.

Other terms and procedures are as explained by Haas and Werner (1969), else as

defined by H. M. Smith (1946 : 17-30) or as presented by Peters (1964).

LACERTILIA

GEKKONIDAE

Hemidactylus turcicus turcicus Linnaeus

Lacerta turcica Linn6, 1758. Syst. Nat., ed. 10, i, p. 202 (Orient).
Hemidactylus turcius, Boettger, 1876. Ber. offenbach. ver. Naturk., 15/16 : 57.
Hemidactylus turcicus turcicus, Mertens, 1925. Abh. senckenberg. naturf. Ges., Frankfurt a.M.,
39 : 60.

MATERIAL EXAMINED (i). BM 1965.683 $, [between Azraq Shishan and Azraq
Druze],* April-May 1965, S. Bisserot.

PHOLIDOSIS. Dorsal tubercles keeled, 14 in a diagonal row across the back.
Usually 2-3 granules between successive or adjacent tubercles. Counts of scansors
are presented in Table i.

MEASUREMENTS. SV : 47 mm ; tail incomplete.

COLORATION. Dorsum with five semi-regular longitudinal rows of brown mark-
ings of irregular shape.

FIELD NOTES. "The only other gecko found was the Turkish Gecko (Hemi-
dactylus turcicus) [one of] which was found in a damaged condition on the track
between Azraq Shishan and Azraq Druze" (S. Bisserot).

REMARKS. Five specimens previously collected in Trans Jordan were available
for comparison (HUJ-R 1063 70 km S. of Amman: Sisah-Chissa, 27 March 1936,

*Not indicated on the specimen label, but derived from Mr. Bisserot's field notes.

220 Y. L. WERNER

Haas; HUJ-R 1064 Hissa-Ma'an, 28 March 1936, Haas; HUJ-R 1068 Wadi Daba'a
SSE of Amman, July 1938, Haas; HUJ-R 1587 Jerash, 18 November 1945, Haas
and Hoofien; HUJ-R 1588 Birketen- Jerash, 16 November 1945, Haas and Hoofien).
Of these, four likewise have 14 rows of tubercles, but one has 12. Scansor counts
for these specimens are included in Table i and compared to Boulenger's (1885) data
for H. turcicus and H. sinaitus. The specimens from Transjordan, including those
from the desert, are typical H. turcicus.

TABLE i

Scansor counts for Hemidactylus turcicus from Transjordan, compared to Boulenger's (1885)
counts for H. turcicus and H. sinaitus. (N, number of specimens.)

Locality (and repository of Scansors (single or paired) under :-

material.) , * ^

Fingers Toes

- %

First

Fourth

First

Fourth

Azraq area (BM)

Jerash area (HUJ)

6-7

9-10

Southern Transjordan (HUJ)

7-8

8-9

6-7

IO-I2

Transjordan, cumulative 6 6-8 8-9 6-7 9-12

Boulenger's H. turcicus 28 6-8 8-10 6-8 9-11

Boulenger's H . sinaitus 157 58

A pattern of dark (brown) spots arranged in 6-8 longitudinal rows occurs in
HUJ-R 1587-8 and, to a lesser extent, HUJ-R 1064. None show a pattern of
crossbands or of large X-shaped designs (each stretched across the back), whereas
these are often encountered in specimens from Cisjordan, the latter pattern being
particularly common in certain localities (e.g. En Gedi).

Ptyodactylus hasselquistii cf. guttatus von Heyden

Ptyodactylus guttatus von Heyden, 1827, in Riippell, Atlas Reise nordl. Afrika, i. Zool.: Rep-

tilien, p. 13, pi. 4, fig. i (Tor, Sinai).
Ptyodactylus hasselquistii Phalanx gutlata (part), Anderson, 1898. Zoology of Egypt, i.

Reptilia and Batrachia, London, pp. 65-67, pi. 6, figs 4, 5 and 5a, pi. 7, figs 6 and 7.
Ptyodactylus hasselquistii guttatus, Barash and Hoofien, 1956. Reptiles of Israel, Tel-Aviv,

p. 161.

MATERIAL EXAMINED (4). BM 1963.664 $ Rum (rocky area), 1963, S. Bisserot.
BM 1965.782 juv. Petra (on red sandstone), 2 August 1965, D. Western. BM
1965.783 and 1965.784 juvs. Wadi Musa (under yellow sandstone rock), 4 August
(3.30 pm and 12.30 pm resp.), D. Western.

PHOLIDOSIS. Tubercles of the three juveniles flat-conical, with indications of
keels ; of the adult, distinctly keeled. Hardly any tubercles in front of the ear or on
the forearm. Other details in Table 2 ("Petra-Rum") which may be compared
with Loveridge's table (1947 : 279).

LIZARDS AND SNAKES FROM TRANSJORDAN 221

MEASUREMENTS. Adult: SV: 68 mm; tail missing. Juveniles: SV: 55, 31 and
30 mm resp. ; only the last with complete (?) tail: 23 mm.

TABLE 2

Pholidosis and measurements of 12 Ptyodactylus hasselquistii sspp. from Transjordan, for
comparison with Loveridge's (1947 : 279) table.

% 6 t % S

DtfltuO 5 3 t"" 1 O I-"

S -3 .S g d * 1 ^

'S 3 'd o fefe ^Sj3

S J2 J9 g g & to

^ O g .5 .5 o Is Is J

Locality (and l| <u ^ ^ ^ g

repository 2 c fc 1 | g 1 J 2 J S

of the material) S fc - | | 1 I -a

* * O, 30 * 38 -3 "3

P i-3 H Q c/) W H H O c/5 G H

Petra - Rum (BM) 4 i 3-4! 11-14 Ir -i3 2 12-13 3~4 9~u 4-6 68 ?

S of Guveira (HUJ) 113 14 11-12 12 2-3 12 6 62-5 54

Basalt Desert (BM) i i 3 12-13 IO 10 2 9 6 41 38-5

Jerash area (HUJ) 6 i 3 10-12 8-10 12 2~5 3 g-n 4 6-7 67 52

1 Four were observed on only one side of one specimen (BM 1965.784).

2 But 15 were counted on one side of one specimen (BM 1963.644).

3 Each of the extreme counts (2 and 5) occurred on only one side of one animal; in all other cases there

were 3-4 scansors under the ist toe.

4 The count is uncertain on one side of HUJ-R 1653 (? 7-9).

COLORATION. Collector's notes: BM 1965.782, "grey with green and brown
spots, light orange mottling". BM 1965.783 and 1965.784, "beige with brown and
white spots". After preservation these three juveniles from near Petra are a light
greyish brown, with small light spots which are nearly round, and are 3-6 granules
across. They are arranged fairly regularly in longitudinal rows ; dark brown spots
of less regular shape are arranged between them. The adult from Rum (BM
1963.664) is coloured similarly, but the light spots are only faintly discernible, and
the dark spots are larger, of more irregular shape, and less regularly scattered. All
specimens have whitish underparts.

REMARKS. Ptyodactylus hasselquistii is notorious for its high geographical vari-
ability (Flower, 1933; Loveridge, 1947; Werner, 1965). This is particularly true in
the regions surrounding the Gulf of Suez, Gulf of 'Aqaba, and the Dead Sea, where
the typical form meets, mixes, or intergrades with guttatus. The present series
shows points of resemblance to guttatus from (central) Israel, and to the original
illustration of von Heyden ; but on the basis of our single adult without tail, alloca-
tion remains uncertain.

One subadult from S of Guveira (HUJ-R 1027, 28 March 1936, Haas) resembles the
present specimens in its keeled dorsal tubercles and in lacking tubercles in front of
the ear and on the forearm. Its particulars are included in Table 2.

222 Y. L. WERNER

Ptyodactylus hasselquistii puiseuxi Boutan

Ptyodactylus puiseuxi Boutan, 1893. Rev. Biol. du Nord de la France, 5 (9) : 27-32, pi. 3, fig. 4

("Bords du lac de Houleh").

Ptyodactylus lobatus syriacus, Peracca 1894. Boll. Mus. Zool. Anat. comp. Torino, 9 (167) :

1-6 (Jerash, Transjordan).

Ptyodactylus hasselquistii puisieuxi, Haas, 1951. Bull. Research Counc. of Israel, i (3) : 95.
Ptyodactylus hasselquistii puiseuxi, Barash and Hoofien, 1956. Reptiles of Israel, pp. 160-161.

MATERIAL EXAMINED (i). BM 1965.682 juv. Basalt desert, Wadi Aseikhim,
April-May 1965, S. Bisserot.

PHOLIDOSIS AND MEASUREMENTS. Tubercles not keeled, each resembling a low
cone. Between ear and corner of mouth 10-15 tubercles, and on each forearm
about 15. Other details in Table 2 ("basalt desert") which may be compared with
Loveridge's table (1947 : 279).

COLORATION. After preservation, dark brownish grey with round whitish dots
(2-3 granules in diameter) alternating with roundish dark spots (5-8 granules in
diameter). Underparts light grey. Tail with conspicuous alternating dark and
light half rings on the dorsal surface ; ventral surface is grey with whitish mottling.

FIELD NOTES. ' . . . the fan-footed gecko (Ptyodactylus hasselquistii) . . .
greater quantities were found when they eventually were observed for the first
time on April 3oth 1965, always on the basalt. One clutch of eleven eggs [obviously
at least 5| clutches] of this species was found at Wadi Aseikhim, nine of which were
hatched and two not. One captive specimen laid two eggs during the journey
back . . . " (S. Bisserot).

REMARKS. The same subspecies occurs at and around Jerash (terra typica,
Peracca, 1894) as also shown by 6 specimens collected by Haas and Hoofien in 1945
(HUJ-R 1651-5 and 6112). Particulars of these are included in Table 2. The
present specimen, closely resembling puiseuxi from Jerash and from northernmost
Cisjordan, allows us to suggest' that puiseuxi is probably primarily associated with
basalt rocks, regardless of whether these are in a mesic habitat (northern Cisjordan
and northwestern Transjordan) or in an arid one (basalt desert of northeastern
Transjordan). It is not, however, absolutely restricted to basalt, occurring also on
adjacent calcareous formations.

AGAMIDAE

Agama pallida haasi subsp. nov.
(Text-fig. 3; Pis i, 2)

Agama ruderata pallida (part), Haas, 1943. Copeia i : 12.

Agama pallida (part), Haas, 1951. Bull. Res. Counc. Israel i (3) : 72-74.

HOLOTYPE. BM 1965.800 $ Azraq in Transjordan, 12 August 1965, D. Western.

PARATYPES (18). Males (10): BM 1965.684 Jebel Uweinid (Basalt desert), April-
May 1965, S. Bisserot; BM 1965.796 Azraq, 12 August 1965, D. Western; HUJ-R
1117 between Sisah and Ma'an, March 1936, G. Haas; HUJ-R 1121 N. Dahaa, 65 m

LIZARDS AND SNAKES FROM TRANSJORDAN 223

SSE Amman, June-July 1938, collector unknown; HUJ-R 1134 between Hissa and
Amman, 28 March 1936, G. Haas; HUJ-R 5215, 5216 and 5217 between Sisah and
Ma'an, March 1936, G. Haas; HUJ-R 1227 near Palmyra, Syria, June 1944, Theodor;
FMNH 48468 Wadi Dabaa 65 m SSE Amman, July 1938, collector unknown (from
Hebrew University). Females (4): HUJ-R 1118 between Hissa and Ma'an, nr.
Ma'an, 28 March 1936, G. Haas; HUJ-R 1120 60 km NE Zerka, Transjordan (no
date), Sjoma Graber; HUJ-R 1124 about 12 km S of Amman, 26 March 1963 (?)
Mendelssohn; HUJ-R 1884, Wadi Debba, Transjordan, Summer 1939, collector
unknown. Juveniles (4): BM 1936.666 Azraq, 16 April 1963, S. Bisserot; BM
1965.797, 1965.798 and 1965.799 Azraq, 12-13 August 1965, D. Western.

DIAGNOSIS. Ear opening distinctly longer than high, bordered above by a row
of conspicuous spines; not round with fairly smooth margin as in Agama pallida
pallida Reuss 1833 from eastern Egypt, Sinai, and southern Israel. Total size
larger, head and body more elongate than in A. p. pallida. Ventral scales usually
smooth, not keeled as in Agama agnetae F. Werner 1939 from western Iraq.
(Pis i & 2).

DESCRIPTION OF HOLOTYPE. A male. Head very convex, short and thick, but
distinctly longer than broad (head index: 114). Nostril not tubular, superior, barely
above the indistinct canthus rostralis. Nasal shield flat. Upper head scales convex,
with short terminal keels ; occipital not enlarged. No well-developed spines on the
hinder part of the head, but a few occipital scales are pointed (resembling the
enlarged scales scattered on the back). A fringe of 3-4 distinct spines on the upper
edge of the ear, pointing downwards (in the preserved specimen), except for one
spine, on the anterior margin of one ear, which points backwards. Ear opening
smaller than eye opening, elongate, nearly twice as long as high; its upper (spiny)
border nearly straight and horizontal (PI. i B). No gular pouch. Body depressed,
not as short as in A. pallida pallida (Pis i A; 2 A). Dorsal scales very small,
irregular, faintly imbricate, indistinctly keeled; intermixed with scattered larger
scales each of which bears a short keel, sometimes ending in a short spine. Scalation
of limbs, and proximal quarter df tail, similar to that of back, but the small ground
scales larger than on back. Ventral scales smooth, imbricate. Tibia longer than
the skull (to occiput). Third ringer shorter than fourth, fifth not extending as far as
second; third toe much shorter than fourth, fifth not extending as far as first. Tail
I 45% SV long (somewhat more than twice as long as the distance from gular fold

MEASUREMENTS OF HOLOTYPE. mm

Total length 189

Snout-vent 77

Head length (to occiput) 20

Head length (total) 24

Width of head 21

Body (occiput-vent) 57

Forelimb 40

Hindlimb 59

Tail 112

224 Y - L. WERNER

to vent), circular in cross section, its distal three quarters with subequal keeled
scales. A double row of 'anal pores' (10 -f 12).

COLORATION OF HOLOTYPE. Collector's note: "mottled dark grey brown: white
dashes". After preservation, brownish grey. Pileus yellowish. In the orbital
area, below the eye, six faint grey radiating streaks; side of head otherwise plain.
Dorsum with four darker brown crossbands, each interrupted by an irregular whitish
vertebral streak. First crossband in front of, second behind, shoulder. Third,
indistinct. Fourth just in front of pelvis. Thirteen uninterrupted crossbands on
tail. The first two are similar in colour to the ones on the body, the remainder are
paler. Underparts light cream, throat mottled with 8-10 grey wavy longitudinal
bands.

VARIATION OF THE TYPE SERIES. Females have no anal pores, and their heads
are shorter (relative to SV length) than in males. The largest specimen is a female
(HUJ-R 1118), SV 93-5 mm; largest male (HUJ-R 1117) SV 89 mm. At the upper
border of the ear there are 2-4 large and 0-2 small spines ; the commonest arrange-
ment is 3 large and i small spine. All juveniles, including the smallest (SV: 33 mm),
show the distinctive ear features of the new form, except that the enlarged scales
bordering the ear opening dorsally are not spiny (PI. i C-D). The ventral scales
are moderately keeled on the posterior abdomen of one specimen (HUJ-R 1134).
The variations of pholidosis, measurements and proportions, and pattern, and com-
parable variations of A. p. pallida from southern Cisjordan are summarized in Table
3. Differences between the samples from Trans Jordan and Cisjordan (comparisons
being made separately among males, females and juveniles), were statistically
significant only in the following instances: Among males, the two samples differed
significantly in SV length (t = 6-32; tio (-05) = 2-23) ; in head length (to occiput) in
%SV (t = 2-59; t 9 (-05) = 2-26); in head width in %SV (t = 4-53; t 28 (-05) = 2-05);
and in the head index (t = 3-52; t28 (-05) = 2-05). Among females the two samples
differed only in head width in %SV (t = 3-18; tzi (-05) 2-08) and in the head
index (t = 2-13; t 2 i ( . 5) = 2-08).

Collector's note on coloration: "The Pale Agamid (Agama pallida) varied con-
siderably, in colour and markings .... A. pallida showed no colour changes
under any circumstances". (S. Bisserot.)

FIELD NOTES. "Three species of ... Agamidae were seen and collected ....
A. pallida was the most common but was found only on the hamada. . . . On
the hamada areas the dominant reptiles appeared to be ... and Agama pall-
ida ... " (S. Bisserot). Two of the adult males (BM 1965.800 and 1965.796)
were caught "among small rough stones: flint, basalt and chert on brown silty
matrix between stones" (the first at 11.30 h). A juvenile, BM 1965.798 "in shade
under Holoxocum silicanum [? Haloxylon salicornicum] near black basalt rocks
(12.30 pm)"; another juvenile, BM 1965.797, "on flint stone desert", and another
among "basalt outcrops large basalt boulders with white interdispersed silt under
rock". HUJ-R 1134 was caught on "Ebene Stein Wiiste" (even stone desert).

"The Pale Agamid . . . appeared to rely on three methods of escape, firstly by
quick bursts of running when the body was held high off the ground and the head up,

LIZARDS AND SNAKES FROM TRANSJORDAN

225

than by flattening the body to the ground and remaining motionless, relying on
camouflage, and lastly by an aggressive stance with mouth open always facing the
attacker." (S. Bisserot.)

GROWTH. The three juveniles collected on 12-13 August 1965 by Mr. Western
measure 44, 38-5 and 33 mm respectively (SV). These obviously had hatched earlier
in the same season. The one taken on 16 April 1963 by Mr. Bisserot measures (SV)
46-5 mm and evidently had hatched in the previous summer. The 17 specimens for
which the date (at least the month) of collection is known (Text-fig. 3) make it
probable that in Transjordan the hatchlings of the year reach ca. 40-50 mm (SV) by
autumn, grow to ca. 70-80 mm during the following year, and attain 80-90 mm in
their third warm season.

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FIG. 3. Agama pallida haasi subsp. nov. Sizes of animals caught at different times of
the year. Squares, males; Circles, females; Diamonds, juveniles. Open symbols,
animals from Transjordan; Solid symbol, animal from near Palmyra, Syria.

REMARKS. It is a pleasure to name this lizard in honour of Professor Georg Haas
who had already commented on its relatively large size (1951). The new taxon is
most closely allied to Agama pallida Reuss 1833 from Eastern Egypt, Sinai and
Southern Cisjordan. In Reuss' original description there is no indication of the
type locality, except that the whole material under discussion had been collected by

226

Y. L. WERNER

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LIZARDS AND SNAKES FROM TRANSJORDAN

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228 Y. L. WERNER

Riippell. Reuss' original specimen label indicates "Aegypt, super." (Klemmer,
1967). The species, however, does not seem to occur in Upper Egypt, and Anderson
(1896 : 79) gives "Sinai" as the type locality (see also Flower, 1933). Through the
courtesy of Dr. Klemmer I could examine excellent photographs of the type (SMF
10007) an d satisfy myself that material from southern Cis Jordan and eastern Egypt
(Kassassin) is in good general agreement with it. Reuss' description includes no
scale counts, but some of his many measurements are represented in Table 3.

The new form is so far known from central and northern Transjordan and SE
Syria, but specimens from the rest of Syria and from Iraq will probably also be found
to belong to this form. A. p. pallida Reuss and A. p. haasi n. ssp. are allopatric,
possibly separated by the steep slopes constituting the eastern rim of the Wadi
'Arava. Though I have seen no intermediate forms, the few specimens from the
Wadi 'Arava being typical pallida, conclusive evidence of reproductive isolation is
not yet available. Thus it seems best to accord the two forms subspecific rank.

In the northern part of its range A. p. haasi may be sympatric with A. ruder ata
Olivier. I have omitted the references for one of these forms having been recorded
within the accepted range of the other, as probably some of these instances are based
on mis-identifications. However, it is my impression that this is not so in all cases
(see also Pasteur and Bons, 1960). It is interesting that it is easier to distinguish
between A. p. haasi and A. ruderata, on the basis of general habitus, than between
A. p. pallida and A. ruderata. Moreover, sexual dimorphism in SV length is
apparently moderate in A. p. haasi (largest $, 89 mm (HUJ-R 1117); largest $,
93-5 mm (HUJ-R 1118)), whereas it is considerable in both A. ruderata (Pasteur and
Bons, 1960) and A. p. pallida (among 117 specimens from Cisjordan in the Hebrew
University collection, largest $, 75 mm (HUJ-R 7509) ; largest $, 87 mm (HUJ-R
5506)). These phenomena apparently represent a case of "sympatric character
divergence" (Mayr, 1965 : 82).

It has been argued that A. pallida Reuss 1833 is conspecific with, and indistin-
guishable from, A. mutabilis Merrem 1820 (Pasteur and Bons, 1960; Wermuth, 1967).
In fact, this possibility had already been mentioned by Anderson (1898). However,
the arguments (and diagrams) of Pasteur and Bons do not entirely exclude the
possibilities that these are either two distinguishable allopatric forms (Flower, 1933)
(with a complex borderline, or intergrading), or even sibling (partly sympatric)
species (Schmidt and Marx, 1956 : 25). It therefore seems most prudent to retain,
at present, the specific name pallida for the populations to which it has traditionally
been applied.

Agama blanfordi fieldi Haas and Werner

(PI. 3 A)

Agama persica fieldi Haas and Werner, 1969. Bull. Mus. Comp. Zool. Harvard, 138 (6) :
337-339, pis. 2-6. (Saudi Arabia: Al-Caissumah - Turaif .)

MATERIAL EXAMINED (i). BM 1965.686 $ Qa el Umari (hard sand desert), 1965,
S. Bisserot.

LIZARDS AND SNAKES FROM TRANSJORDAN 229

PHOLIDOSIS. Dorsal scales subequal, keeled and shortly mucronate. Lateral
scales similar but smaller. Ventral scales feebly keeled. Scales around middle of
body, 80.

MEASUREMENTS. SV, 105 mm; tail, 163 mm.

COLORATION. Collector's note: "A. persica when first captured turned to a
brilliant blue in the area of the dew-lap under the chin but not on any other part of
the body. Both A. sinaita and A. persica turned blue when killed and preserved in
spirit." (S. Bisserot.) Yellowish grey*. Dorsally, four darker longitudinal bands,
brown with still darker margins. Each band is of uneven width, and contains about
six alternating dilated and constricted zones. Through the dilated zones run trans-
verse series of white dots which are interrupted by a median light band. Each dot
coincides with an enlarged scale, making it more conspicuous. On the head there
are two brown wavy crossbands, preceded by a longitudinal patch of the same colour.
On the tail the dorsal pattern gradually changes to one of simple dark rings. Belly
with a distinct central longitudinal, grey band, and irregular lateral ones. Gular
pouch dark grey with remnants of blue.

FIELD NOTES. "Three species of ... Agamidae were seen and collected . . .
on the flat dried sand areas only the one specimen of A. persica was found". S.
Bisserot.

REMARKS. This specimen is identifiable according to Boulenger's (1885) key as
A. blanfordi (S. C. Anderson, I966a; nom. subs, for A. persica Blanford 1881, nom.
preoccup.) because of its unequal dorsal scales. However, the gular pouch and the
scattered enlarged scales are less developed than in blanfordi and the head and body,
especially the former, are more depressed than in this form. On the other hand, a
series of very similar specimens collected in NE Saudi Arabia by Mr. Henry Field
shows considerable variation in the development of the enlarged dorsal tubercles;
several specimens have homogenous scaling and are thus identifiable as isolepis
(Boulenger, 1885, lectotype from between Magas and Bampur, southeastern Iran
S. C. Anderson, i966b). Similar, apparently, were the two specimens from Meso-
potamia which Steindachner (1917) identified as "isolepis with a unique pattern".
Our specimen is very similar to the pair depicted by him (at least in proportions,
pattern, and non-meristic scale characters).

Apparently this is a form allied both to the agilis-isolepis Rassenkreis (Wettstein,
1951) and to blanfordi. It is, in certain respects, intermediate between the two. It
is characterized by variably (mostly feebly) developed dorsal tubercles and gular
pouch, and by a very distinctive pattern of longitudinal bands. A confusion con-
cerning agilis and blanfordi had already been suspected by Schmidt (1941).

This new form has previously been referred to as A. persica Blanford. Thus at
least part of the series mentioned by Haas (1957) belongs to A. blanfordi fieldi, as
judged by specimen CAS 84541 (now HUJ-R 7081) and by his description of the
pattern of CAS 84477. The latter description has been accepted by Khalaf (1959)

*When examined by the author.

230 Y. L. WERNER

as applying to A. persica Blanford though differing from that usually encountered
in this species.

Agama sinaita von Heyden

(PI. 3 B-E)
Agama sinaita von Heyden, 1827. In Riippell, Atlas Reise nord. Afrika, Kept., p. 10, pi. 3 (Sinai).

MATERIAL EXAMINED (4). BM 1965.685 <$ Wadi Ratan [W. Ratam] (basalt
desert), April-May 1965, S. Bisserot. BM 1965.801 <f>, BM 1965.802 $ Petra (on red
and yellow sandstone respectively), 2 August 1965, D. Western. BM 1965.803 $
Wadi Musa, near Petra (basking on soil on top of rock), 4 August 1965, D Western.

PHOLIDOSIS. In the specimen from the basalt desert (BM 1965.685) the tail is
moderately compressed laterally including its thick basal portion. The two dorsal
rows of caudal scales are enlarged, their thick keels creating the impression of a
slight crest. The specimens from Petra and Wadi Musa resemble specimens from
Cisjordan in that the thick part of the tail is nearly cylindrical, and carries dorsally
four straight rows of strongly keeled scales, the two median rows being little better
developed than their immediate neighbours (PI. 3 C & E).

MEASUREMENTS. SV: 103, 69, 79, 79 mm. Tail of the last: 125 mm (other tails
incomplete). In all specimens the third toe is hardly longer than the fourth (PI. 3 B).

COLORATION. "The ability of A. sinaita to change colour was observed on several
occasions but did not appear to follow a regular pattern. Most specimens were
observed to be a brilliant cobalt blue on first sight [Mountfort, 1965 : plate 4oa] but
changed to a dark chocolate brown when pursued. One specimen kept alive changed
from brown to blue over the head and shoulders and part of the flanks when food was
put in its mouth. This reaction was repeated in captivity in this country [England]
and was also caused by the temperature being raised to 80 F [27 C] or higher . . .
A. sinaita turned blue when killed and preserved in spirit." (S. Bisserot.)

Professor H. Mendelssohn (Tel- Aviv University) has studied the colour changes of
this species, as part of his research of its behaviour, and found marked sexual
dichromatism. Hence the arrangement of the following notes on our specimens.

Males: BM 1965.865 (preserved) : blueish grey (throat and belly darker), tail grey
(yellowish ventrally). 1965.802, collector's note: "bright blue all over, faded after
death." The preserved specimen is dark grey, nearly black, with underparts lighter
(and posteriorly very light) brownish grey.

Females'. Collector's notes summarized: "head blue when alive, turning brighter
blue when killed. Body grey with orange blotches". After preservation the heads
are blackish, the bodies dark grey, and the tails have alternating darker and lighter
transverse bands. The throats are grey (reticulated in 1965.801) and the abdomens
steel grey; the remaining underparts are cream- white.

REMARKS. The unusually large male with compressed tail, from the basalt
desert, conceivably represents a distinct subspecies, but this cannot be assessed on
the basis of a single specimen. (Wettstein, 1951 : 433, mentions three specimens

LIZARDS AND SNAKES FROM TRANSJORDAN 231

from northern Trans Jordan, but gives no particulars apart from the colour.) The
remaining specimens have tails similar to those of animals from southeastern Israel
and northeastern Sinai, and the same is true of two collected by Haas between
Guweira and Aqaba in 1936 (HUJ-R 1137, HUJ-R 5231). On the other hand, all
the specimens from Trans] ordan are characterized by the third toe being hardly
longer than the fourth. Specimens from southeastern Israel and northeastern Sinai
usually have a longer 3rd toe (PI. 3 B & D).

Agama stellio brachydactyla Haas
(Text-figs 4, 5)

Agama stellio brachydactyla Haas, 1951. Ann. Mag. Nat. Hist. (Ser. 12), 4 : 1052 (Israel : foot
of Jebel Lussan, near Israel-Sinai frontier, S.S.W. of Beer-Sheba).

MATERIAL EXAMINED (3). BM 1965.787-789, Basta (remarks on habitats, under
coloration), 3 August 1965, D. Western.

PHOLIDOSIS. A mid-dorsal band of unequal enlarged scales, about six times as
broad as one of the larger scales. The transverse series of tubercles extend across
this band. They are slightly interrupted medially but here some of the interstitial
scales are almost as large as the tubercles themselves. All large scales are either
distinctly keeled, mucronate, or spinous (Text-fig. 4).

FIG. 4. Agama stellio brachydactyla. Mid-dorsal sclaes of BM 1965.787 from southern
Transjordan (Basta). (From a photograph.) Scale, 10 mm.

Lamellae under ist finger: 9, 10, 9. Under 4th toe: 22, 24, 22.

MEASUREMENTS. SV: no, 84, 103 mm. Tail of smallest: 116 mm; other tails
incomplete. Foot hardly longer than tibia, being shortest in largest specimen.

COLORATION. Collectors notes: 1965.787, "Back: black; black and yellow trans-
versely striped tail; orange blotches near head (basking on orange mauve quarzite,

232 Y. L. WERNER

3.25 pm)." 1965.788, "Yellow and black with orange spots on neck. (On yellow
sandstone, 4.20 pm)". 1965.789, "Brownish with orange and brown blotches.
Tail black with orange transverse stripes. (On soil besides flintstone, 3 pm)".

In the preserved condition all three specimens are grey dorsally with pale yellowish
blotches, the largest blotches arranged in mid-dorsal asymmetrical pairs, each pair
tending to fuse and to form a large obliquely transverse blotch. On the tail, trans-
verse bands of the same yellowish colour, which also covers the underparts, the
throat being faintly reticulated with pale grey.

REMARKS. All three specimens are presumably not fully grown. These indi-
viduals are not very typical brachydadyla. In the number of lamellae under the
toes, as well as in the relative size of the mid-dorsal scales, they rather resemble
specimens from the northern Negev in Israel, which are intermediate between
brachydadyla and the form inhabiting mediterranean Israel. The specimens from
Basta are assigned here to brachydadyla in accordance with their coloration and also
in order to indicate their geographical affinities (Daan, 1967).

Six other specimens from Transjordan are available. Only one from Petra
(HUJ-R 1096, 29 March 1935, Haas) shows a similar arrangement of a mid-dorsal
band of subequal enlarged scales. Two other specimens from southern Transjordan
(HUJ-R 1094 and 1103) and three from the Jerash-Amman area (HUJ-R noi,
1 1 10 A, and mo B) have the dorsal transverse series of tubercles clearly separated
by smaller scales, as is usual in specimens from northern Cisjordan.

The number of lamellae under the 4th toe in the nine specimens from the two areas
in Transjordan is presented in Text-fig. 5 which also includes, for comparison,
samples from five localities in Cisjordan. As the figure shows, in Cisjordan there is
a pronounced north-south gradient in this character (with the higher values in the
north). A parallel but less prominent gradient is indicated in Transjordan.

CHAMAELEONIDAE

Chamaeleo chamaeleon recticrista Boettger

Lacerta chamaeleon Linnaeus, 1758. Syst. Nat., p. 204 (Africa and Asia.)

Chamaelo vulgaris var. recticrista Boettger, 1880. Jahresber. senckenberg. naturf. Ges.
Frankfurt, p. 198 (Jeruslaem and Haifa).

MATERIAL EXAMINED (i). BM 1963.667 juv. between Shaubak and Tafila (on
camel thorn), 1963, S. Bisserdt.

FIG 5. Agama stellio subspp. Numbers of lamellae under fourth toe in Trans- and Cisjor-
dan. Material included in each sample was collected within the area of the map covered by
the relevant circle. (A) Jerash-Amman area; (B) Desert locality included in sample C;
(C) Petra-Basta area; (D) Hills surrounding Lake Tiberias; (E) Ramot-HaShavim ;
(F) Jerusalem and adjacent Judaean Hills; (G) Be'er-Sheva and vicinity; (H) Sde-
Boker-'Avdat area; M, Mean; N, Number of specimens. (Political frontiers as before
1966.)

LIZARDS AND SNAKES FROM TRANSJORDAN

233

234 Y - L. WERNER

PHOLIDOSIS. Gular and abdominal crest of enlarged scales present.
MEASUREMENTS. SV: 44 mm; tail: 42 mm.

COLORATION. Grey with irregular dots of darker grey. On each flank two longi-
tudinal rows of five light cream blotches. Enlarged scales of dorsal, gular and
abdominal crests the same light cream.

REMARKS. A half -grown specimen from El-Hamma (NW Transjordan, in Israel
HUJ-R 1501; 13 March 1945, Coll. G. Haas) is similarly coloured. In a juvenile
from Jerash (HUJ-R 1502; 15 November 1945, Coll. Haas and Hoofien), measuring
36 mm (SV) the occipital casque is not (yet) developed. All three specimens conform
to the chamaeleons of northern Israel rather than to the C. c. musae-like animals
from further south (Hoofien, 1964).

LACERTIDAE

Acanthodactylus boskianus asper Audouin

Lacerta aspera Audouin, 1829. Descr. Egypte, Kept., Suppl., p. 173, pi. i fig. 9 (Egypt).
Acanthodactylus boskianus var. asper Lataste, 1885. Ann. Mus. Geneva 2 (2) : 496.

MATERIAL EXAMINED (3). BM 1965.691 $ and BM 1965.693 $ Tell Quarma
[=Tell Qarma] (blown sand wadi), April-May 1965, S. Bisserot. BM 1965.804 $
Wadi Musa near Petra (sandy soil near bushes), 4 August 1965, D. Western.

PHOLIDOSIS. Scales across middle of body: 31, 38, but 55 in BM 1965.693.
Gular scales in straight median series: 26, 32, 26. Lamellae under 4th toe: 21-22.

MEASUREMENTS. BM 1965.691, 56 mm SV, 121 mm tail. BM 1965.693, 81 mm
SV, and BM 1965.804, 62 mm SV (tails incomplete).

COLORATION. BM 1965.804, collectors note: "brown: long orange stripes,
spotted with black".

REMARKS. Eleven other specimens from various localities in southern Trans-
Jordan (HUJ-R 1335, 1338, 1341, 1661, 5041-6, 5053) have the following pholidotic
counts: Scales across middle of back, 29-51 (against 29-42 in southern Cisjordan,
N = 24). Gular scales in a straight median line, 25-35 (24-31, in Cisjordan,
N =24). Lamellae under 4th toe, 19-22 (as in Cisjordan). Other conventional
counts are also similar in Cis- and Transjordan except that some Cisjordan specimens
show a reduction of the lateralmost ventral plates, so that only 8-9 longitudinal
series are present.

In general, adult male A. b. asper are larger than females. The 2 largest males
seen from Transjordan are 75 and 77 mm (SV), so that the female from Tel-Qarma
(81 mm) appears unusually large.

The dorsal pattern of some Transjordan males includes rows of sharply defined
blackish dots, instead of the more usual rows of irregular brownish spots.

LIZARDS AND SNAKES FROM TRANSJORDAN 235

Acanthodactylus grandis Boulenger
(PI. 4 A, B)

Acanthodactylus grandis Boulenger, 1909. Ann. Mag. Nat. Hist., 4 (8) : 189 (Jerud and Ataiba,
Syria) .

MATERIAL EXAMINED (2). BM 1965.692 $ Ain el-Enoquiyya (sand and stone
wadi) ; BM 1965.694 $ Tell el Mukheizin (Hamada, beneath carcass of dog) ; both
April-May 1965, S. Bisserot.

PHOLIDOSIS. Scales across middle of body: 60; 60. In BM 1965.692 there is a
fifth small upper labial before the center of the eye, on each side.

MEASUREMENTS. BM 1965.692: SV: 73 mm; tail: 126 mm. BM 1965.694:
SV: 96 mm (tail incomplete).

COLORATION. BM 1965.692 (PI. 4 B) : Black spots, each covering up to ten scales,
arranged in ten regular longitudinal rows, and in irregular transverse series. The
two median rows begin at the occiput but disappear before the middle of the back ;
4th and 5th row on each side present on flanks but absent from neck. Some rows
extend on the tail, on the lateral sides of which the spots are represented as vertical
blotches at every second suture between scale rings. Ground colour (preserved),
nearly uniform grey (compare PI. VI of Boulenger, 1923).

BM 1965.694 (PI. 4 A) : Six dark longitudinal stripes faintly indicated on back.
Along these there are a few, irregularly scattered, small blackish spots, each covering
up to 7 scales.

In both specimens the sides of the head bear alternate light and dark vertical bars,
one of the latter passes through the eye.

REMARKS. All characters of both specimens are within the range of variation
shown by A . grandis in the HU J collection, some of which have been mentioned by
Haas (1943; Transjordan between Hissa and Ma'an). There is some difficulty in
distinguishing immature A. grandis from A. scutellatus scutellatus, which likewise
has smooth scales, since the range of variation of almost all conventional scale
counts is nearly identical (see also Boulenger, 1923 : 50). BM 1965.692 resembles
A. s. scutellatus in the number of supralabials (5) before the center of the eye, and in
its relatively long foot with moderately well developed pectination. The specimen
however is certainly assignable to A . grandis for the following reasons :

The snout with its somewhat swollen nasals resembles that of other A . grandis, and
not at all the pointed snout of A . s. scutellatus.

While five supralabials in front of the center of the eye are characteristic of A. s.
scutellatus, and 4 of A. grandis, 5 may sometimes occur in the latter (see also
Boulenger, 1921 : 114-115).

The longer foot and relatively stronger pectination (when compared to large A.
grandis such as BM 1965.694) appear to be largely juvenile characters, which are
paralleled in series of other species of Acanthodactylus containing mature and
immature specimens. Moreover the pectination is still far less developed than in
adult A. s. scutellatus from southern Israel.

236 Y. L. WERNER

The pattern conforms closely to that of A. grandis and differs most strikingly
from that of A . s. scutellatus, as the latter never show any longitudinal arrangement
of the markings.

Acanthodactylus tristrami tristrami Gunther
(PI. 4 C, D)

Zootoca tristrami Gunther, 1864. Proc. Zool. Soc., p. 491 (Lebanon).
Acanthodactylus tristrami Boulenger, 1881. Proc. Zool. Soc., p. 746, pi. 64, fig. i.

MATERIAL EXAMINED (i). BM 1962.352 <j> Jordanian-Syrian border, Jerusalem-
Damascus Rd (Outside customs shed) [probably loc. i on map], 4 June 1952, W.
Larmuth.

PHOLIDOSIS. Scales across middle of body: 59. Longitudinal rows of ventrals:
ii. Other characters also in agreement with Boulenger's (1921) data for A.
tristrami.

MEASUREMENTS. SV: 82 mm; tail (tip missing) : 86 mm.

COLORATION. The blackish markings tend to form a reticulum along each side
of the dorsum (PI. 4 C).

FIELD NOTES. "Died while ovipositing".

REMARKS. Angel (1936) described from NE Syria A. t. orientalis (48-56 scales
across middle of body), which was also reported from the neighbourhoods of Rutba
(Schmidt, 1939) and Mosul (Haas, 1952) in Iraq. From Haditha, Iraq, Schmidt
(1939) described A. t. iracensis (45-46 scales across middle of body). Thus
Giinther's (1864) and Boulenger's (1921) A. tristrami was accorded subspecific rank
as A. t. tristrami (58-65 scales across middle of body).

Our specimen appears assignable to the typical form, as well as two specimens in
the HUJ collection, reported by Haas (1943) : HUJ-R 1333 <$, 15 km S of Amman,
SV 92 mm; HUJ-R 1332 <j>, 45 km S of Amman, SV 69 mm (after a year in captivity).
Scales across middle of body, 58, 57 respectively. Ventrals in 10 rows.

All three specimens are larger than Angel's A. t. orientalis (1936; 50-66 mm snout-
vent, N = 8), although this alone would not have been taxonomically significant.

The dorsal pattern of HUJ-R 1333 (<) consists of distinctly X-shaped blackish
marks (PI. 4 D; Boulenger, 1921). Markings intermediate between this pattern and
the reticulum of BM 1962.352 (?) are present on the female (the type ?) figured by
Tristram (1885: PI. 16, fig. 2).

Eremias brevirostris microlepis Angel

Eremias brevirostris microlepis Angel, 1936. Bull. Inst. Egypte 38 : 112-113 ("Haouarine"
5 5 km SE of Horns, Syria) .

MATERIAL EXAMINED (2). BM 1965.689 $ (?) Qasr Amra (hamada) ; BM 1965.690
$ (?) Shishan (hamada) ; both, April-May 1965, S. Bisserot.

LIZARDS AND SNAKES FROM TRANSJORDAN 237

PHOLIDOSIS. Scales across middle of body: 60; 62. Longitudinal series of
ventral plates: 10. Plates in collar: 9. Gular scales in a straight median series: 29;
28. Femoral pores: 15, 14-15. Lamellae under 4th toe: 20; 24. Upper labials
anterior to centre of eye: 5, the 5th being the first of two small false supralabials
below the subocular.

MEASUREMENTS. SV 46; 49 mm. Tail : 69 (tip missing) ; 81 mm.
COLORATION. Both pale, the ocelli inconspicuous.

FIELD NOTES. "The blown sand areas in wadis were the chief habitat of the
fringe-toed lizards, Acanthodactylus . . . however the lizard (Eremias brevirostris)
was also seen in this habitat but not as frequently as on the hamada" (S. Bisserot).

REMARKS. Specimens from eastern and north-eastern Syria, like those from
Iraq, are so far inseparable from the typical form (Angel, 1936 ; Schmidt, 1939 ; Haas
and Werner, 1969). Angel's microlepis from western Syria (and a greater altitude)
had been based on a single specimen, and was not regarded as valid by Haas (1957 :
73). However, the present two specimens from northern Trans Jordan agree fairly
well with Angel's description. Furthermore, 13 specimens from central Trans-
jordan (Amman-Ma'an) in the collection of the Hebrew University (Haas, 1943 : 14)
show a clear affinity to microlepis, having 46-57 (commonest numbers 53-54) scales
across the middle of the body, and also relatively small gular scales (21-29, usually
25-27, in a straight series). Interestingly a specimen from southern Trans Jordan
(Guweira-Aqaba, HUJ-R 1230) has only 44 scales across the middle of the body (and
25 gulars). For comparison, Angel's (1936 : 112) E. b. brevirostris from NE Syria
had 40-52 (commonest numbers 47-49) scales across the middle of the body, and
20-25, usually 21-23, gular scales in a straight series. Thus E. b. microlepis occupies
the centre (around Jebel ed Druze) of the western distributional frontier of the species
(Hoofien, 1957), possibly intergrading with the typical form to the north, east and
south.

Eremias guttulata guttulata Lichtenstein

Lacerta guttulata Lichtenstein, 1823. Verz. Doubl. Mus. Berl., p. 101 (Egypt).

Eremias guttulata, A. Smith, 1845. 111. Zool.. S. Afr., Kept., PI. 48, fig. 8.

Eremias guttulata forma typica, Boulenger, 1921. Monograph of the Lacertidae, London, 2,

p. 258.
Eremias guttulata guttulata, Wettstein, 1928. Sitzber. Akad. Wiss. Wien (math.-natur.) 137,

Abt. i, p. 782.

MATERIAL EXAMINED (i). BM 1965.688 $ 2 miles S of Azraq Druze (basalt
desert), April-May 1965, S. Bisserot.

PHOLIDOSIS. Scales across middle of body: 50. Longitudinal series of ventral
plates : 8. Femoral pores : 13-14. Lamellae under 4th toe : 22-24. Upper labials
preceding subocular (which enters lip) : 4.

MEASUREMENTS. SV: 46 mm. Tail: 83 mm (tip regenerated).

238 Y. L. WERNER

COLORATION. The dark borders of the dorsal 'ocelli' are black and tend to merge
with their neighbours laterally, forming incomplete black crossbands, which are
interrupted by the white centres of the 'ocelli'.

REMARKS. The snout is very elongated, pointed and flattened. In comparison
with specimens from Cis Jordan the pileus is smooth and flat, eyes and nostrils being
little elevated.

Fourteen other specimens from various localities between Amman and Petra
(HUJ-R 1237, 1240, 1256-7, 1259-60, 1262-3, 6236-8, 6273-4, 6300) have the
following ranges of counts: Scales across middle of body, 44-57 (48-52 in 9 speci-
mens). Longitudinal series of ventral plates, 10. Femoral pores, 10-14 ( I2 ~ I 3 m
9 specimens). Lamellae under 4th toe, 18-24 ( 22 m 6 specimens). Upper labials
preceding subocular, 4. The pileus, as in the Basalt Desert specimen, is relatively
smooth and flat, although in a few specimens the nostrils (and sometimes the eyes
too) are somewhat elevated.

The 3 largest specimens measure 50-51 mm (SV).

In none of these 14 specimens does the dark component of the pattern occupy
such a large area as in the specimen from the Basalt Desert, nor is this component
black. In some specimens it is brown, in others, pale to the point of becoming
indistinct. Some specimens show a tendency for lateral confluence of 'ocelli', but
the resulting pattern resembles strings of beads rather than crossbands of uniform
width. The range of coloration known from Cisjordan resembles that shown by
these 14 specimens. Thus the Basalt Desert specimen is outstanding in its black
and extensive dark pattern, perhaps as an adaptation to its habitat.

Ophisops elegans blanfordi Schmidt
(Text-figs 6, 7)

Ophisops blanfordi Schmidt, 1939. Zool. Ser. Field Mus. Nat. Hist., 24 (7) : 64-65 (Halfaya,
20 miles east of Amara, Iraq).

MATERIAL EXAMINED (2). BM 1963.668 ^ N of Shaubak, 1963, S. Bisserot; BM
1965.687 $ Ain el Enoquiya (sand and stone wadi), April-May 1965, S. Bisserot.

PHOLIDOSIS. Scales and plates around middle of body 38; 34. Femoral pores:
10-11 ; 10-11. Lamellae under 4th toe : 22-24. Upper labials preceding subocular:
4. Third postsubocular in broad contact with auricular. Postnasal: single.
Occipital of medium size (somewhat larger than postnasal).

MEASUREMENTS. SV: 42; 40 mm. Tail of <$: 92 mm; of $ missing.

COLORATION. Both have the usual Ophisops elegans pattern except that there is
a distinct dark vertebral line running from the occiput to the pelvic region.

REMARKS. Both specimens agree with Schmidt's description, except in having a
slightly higher number of scales around the body (Schmidt's 92 specimens, all from
the lower Tigris-Euphrates Valley, had 30-36 scales and plates around the middle of
the body, averaging 33).

LIZARDS AND SNAKES FROM TRANSJORDAN 239

The dark vertebral line observed in the two specimens from Transjordan occurs
only rarely in 0. e. ehrenbergi from Cisjordan (N = 50), and then only on the neck.
Likewise, Lantz (1930 : 41) says of the pattern of 0. e. elegans "Dessin caracterise
per la bande occipitale rudimentaire . . . Bande occipitale absente ou reduite a un
petit trait ou a quelques petites taches noires sur la nuque." It is much commoner,
and better developed, in 0. e. schlueteri from Cyprus (N = 16).

Ophisops from Transjordan in the HUJ collection fall into two groups. All
those from Jerash and its vicinity (N =16: HUJ-R 1190, 1203, 1204/1-2, 1205,
1206/1-4, 1207/1-3, 1208/1-2, 1209, 1561) have double postnasals. In most of
them, the vertebral line is either absent, or confined to the occipital region; but in
3 $3 it extends to the shoulders and in a single male it reaches the midbody although
it is very faint. These specimens appear to be assignable, like all those from Cis-
jordan, to 0. e. ehrenbergi. On the other hand, specimens from between Amman
and Petra are assignable to blanfordi (N =9: HUJ-R 1183, n86, 1218, 1220-22,
6158-60). Of these, 5 have single postnasals, 3 have double postnasals, and one is

i/>

H-
I/)

O n D..O D cm

oo n v o DO on

1/2

01 23456

VERTEBRAL LINE

FIG. 6. Ophisops elegans subspp. from. Transjordan. Number of postnasals against extent
of vertebral line (o, none; 2, only on occiput; 3, reaching shoulder; 4, reaching midbody;
5, reaching pelvis; 6, reaching tail base. These values are adjusted by + i for unusually
intense lines, and by -i for particularly faint ones). Open symbols, O.e. ehrenbergi from
the Jerash area; Solid symbols, O. e. blanfordi (details in text); Squares, males; Circles,
females; Diamonds, juveniles.

240

Y. L. WERNER

asymmetrical. In most the vertebral line is well developed and this is particularly
true of those with double postnasals. Thus the two forms are distinguishable by the
combination of these two characters (Text-fig. 6). Possibly they are also distin-
guishable by a combination of femoral pore number and vertebral line extent

(Text-fig. 7).

a 13

a.
to

oe.

1 1

o 10

D
D

a
oo

0123456

VERTEBRAL LINE

FIG. 7. Ophisops elegans subspp. from Transjordan. Number of femoral pores (repre-
sented for each specimen as the mean of both femurs) against extent of vertebral line.
Symbols as in fig. 6.

The 9 HU J blanfordi specimens have the following scale counts : Scales and plates
around middle of body, 29-44 (39-41 in 4 specimens) ; Femoral pores, 8-13 ; Lamellae
under 4th toe, 20-25. Neither these nor the remaining conventional counts differ
markedly from those found in 0. e. ehrenbergi.

In conclusion, while the Ophisops of the Jerash district appears to be consub-
specific with 0. e. ehrenbergi of Cisjordan, the form occurring in the more arid parts
of Transjordan is Schmidt's 0. blanfordi. This, however, does not seem to merit
specific rank since there are indications of intergradation with 0. elegans in Trans-
Jordan (details above) and Iraq (Haas and Werner, 1969), and there is no evidence of

LIZARDS AND SNAKES FROM TRANSJORDAN 241

sympatry. Two additional alleged blanfordi characters in fact occur also in (other)
0. elegans: the small temporal scales, which occur in 0. e. schlueteri (Cyprus) ; and the
situation of the third post subocular which in 0. e. ehrenbergi (Cisjordan) and 0. e.
schlueteri sometimes touches and sometimes fails to touch the auricular.

SCINCIDAE

Chalcides oceltatus ocellatus Forskal

Lacerta ocellata Forskal, 1775. Descr. Anim., p. 13 (Egypt).

Chalcides ocellatus forma typica, Boulenger, 1890. Ann. Mag. Nat. Hist. 5 : 444-445.
Chalcides ocellatus ocellatus Wettstein, 1928. Sitzber. Akad. Wiss. Wien (math.-natur.), 137,
Abt. I, p. 784.

MATERIAL EXAMINED (i). BM 1965.785 Aqaba (on rocks close to beach, Red Sea
coast), 5 August 1965, D. Western.

PHOLIDOSIS. Scales around the middle of the body: 30.
MEASUREMENTS. SV 61 mm; tail: 69 mm.

COLORATION. Collector's note: "brown with darker brown and white spots".
The 'ocelli' are numerous and are arranged in transverse series on the neck and tail.
They are small, each occupying less than a single scale.

REMARKS. Two out of 3 specimens (the 3rd being damaged) from 65 miles SSE of
Amman (HUJ-R 1442, 5107, 5108 June-July, 1938 Haas) have 28 scale rows.
The same applies to HUJ-R 1469 from El-Hamma (NW Transjordan within Israel
13 March 1945, Haas). The pattern of HUJ-R 5107 is remarkably irregular, many
of the 'ocelli' having only a black spot on one side of the white centre instead of a
complete border. In HUJ-R 1442 and 5108 (juveniles) there are only faint indica-
tions of 'ocelli'. The specimen from El-Hamma has 'normal' 'ocelli', each occupying
a scale.

Eumeces schneideri princeps Eichwald
(PL 5 B)

Euprepes princeps Eichwald, 1839. Bull. Soc. Imp. Nat. Moscow, 2 : 303-307 ("In ora Caspia

occidentali, ad montes praesertim Talyschensis") .
Eumeces princeps, Taylor, 1935. Kansas Univ. Sci. Bull., 23 : 138.
Eumeces schneideri princeps, Eiselt, 1940. Zool. Anz., 131 : 218.

MATERIAL EXAMINED (i). BM 1965.695 <$ Ain el Enoquiyya (basalt desert),
April-May 1965, S. Bisserot.

PHOLIDOSIS. Scales around middle of body: 27. Dorsal scales from occiput to
above cloaca: 67.

MEASUREMENTS. SV: 115 mm (tail regenerated).

COLORATION. Ground colour of back light brown. No light spots. Along the

242 Y. L. WERNER

flank a dark brown band, 2-3 scales broad; its lower border half a scale above the
light lateral band, its upper border fairly sharp (PI. 5 B). The light lateral band is
intensely white. (It may have been yellow at the time the animal was killed, six
months prior to its examination by me.)

REMARKS. The scale counts of this specimen are characteristic of princeps
(Eiselt, 1940: Table i), but the colour is unusual, and may represent an adaptation
to the basalt desert. A specimen from Shaubak (BM 1963.669) here assigned to
schneideri shows some tendency towards a similar coloration. However, a specimen
collected 65 miles SSE of Amman (HUJ-R 1389) and clearly assignable to princeps
(27 scale rows, 69 dorsal scales from occiput to above cloaca) is uniformly coloured
having neither light spots nor darkened flanks, the light lateral band merging with
the light belly.

Eumeces schneideri schneideri Daudin
(PI. 5 A, C)

Scincus schneideri, Daudin, 1802. Hist. Nat. Rept., 4 : 291.

Eumeces schneideri (part*), Taylor, 1935. Kansas Univ. Sci. Bull. 23 : 126.

Eumeces schneideri schneideri, Eiselt, 1940. Zool. Anz., 131 : 213.

MATERIAL EXAMINED (2). BM 1963.669 N of Shaubak, 1963, S. Bisserot. BM
1965.786 Petra (on red sandstone), 1965, D. Western.

PHOLIDOSIS. Scales around middle of body: 26, 24. Dorsal scales from occiput
to above cloaca : 65, 66.

MEASUREMENTS. Largest (BM 1965.786): SV: 114 mm; tail: 209 mm.

COLORATION. Both specimens have the usual light lateral band passing through
the ear. BM 1963.669 (PI. 5-C) : Only a few small light (originally yellow-orange ?)
spots, each covering up to a third of a scale. Flanks mottled dark brown above the
light band; each dark spot covering the posterior portion of a scale. BM 1965.786
(PI. 5 A): collector's note: "green with orange spots". The spots each cover up to
a whole scale, and are irregularly arranged; a tendency to form transverse series is
particularly evident on the tail.

REMARKS. These specimens were collected relatively near the area where, in
Israel, the northwestern pavimentatus, and the southern schneideri intergrade. In
southern Israel and in Sinai (Schmidt and Marx, 1956 : 28) there occur populations
in which the pattern is regularly of the schneideri type, but the scale counts of many
specimens tend towards those characterizing pavimentatus (Eiselt, 1940: Table i).
This situation is exemplified by BM 1965.786 from Petra, which it seems best to
assign to schneideri (see also Taylor, 1935 : 130) like the specimens from southern-

*Taylor includes in schneideri single specimens from "Haiffa" and "Mt. Jerusalem", within the range
of pavimentatus. These specimens evidently are adult males of pavimentatus which, unlike the females,
lose the whitish streaks adorning the young. The orange spots however, remain arranged in longi-
tudinal rows (see his Plate 5).

LIZARDS AND SNAKES FROM TRANSJORDAN 243

most Israel. BM 1963.669 from North of Shaubak has 26 scale rows, and its very
broad dorsal scales exclude it from princeps.

Mabuya vittata Olivier

Scincus vittatus Olivier, 1804. Voy. Emp. Ottoman, 3, p. 103, pi. 29, fig. i (sands west of

Rosetta) .
Mabuia vittata Boulenger, 1887. Cat. Lizards Brit. Mus., 3, p. 176.

MATERIAL EXAMINED (i). BM 1963.670 2 km SE Druze village, Azraq, 1963,
S. Bisserot.

PHOLIDOSIS. Scales around middle of body: 32.
MEASUREMENTS. SV: 73 mm; tail: 80 mm.

COLORATION. Dorsum brown, with three light longitudinal bands. No darker
spots, except tiny ones on occiput.

REMARKS. Of 4 specimens from the surroundings of Jerash (HUJ-R 1423, 1424,
1426, 1547; November 1945, Coll. Haas and Hoofien), 3 have 32 scales around the
middle of the body, and one has 34. The largest of these measures 95 mm (SV).
The pattern varies: one specimen resembles BM 1963.670, but another has 5 light
bands, and two have 4, the median one being obliterated. The other 3 specimens
have most dorsal scales partly edged in black (or dark brown), particularly towards
the borders of the light bands. In Cisjordan, too, the pattern and colour of this
species are highly variable (cf. Peracca, 1894 : 8).

OPHIDIA
COLUBRIDAE

Natrix tessellata tessellata Laurent!

Coronella tessellata Laurenti, 1768. Synops. Kept.: 87 ("in Japidia, vulgo Cars").
Natrix tessellata, Bonaparte, 1834. Iconogr. Faun. Ital., 2, n : plate.
Natrix tessellata tessellata, (Hecht) 1930. Mitt. zool. Mus. Berlin, 16 : 319.

MATERIAL EXAMINED (i). BM 1965.696 juv., Shishan (sandy area nr. date palms),
1965, S. Bisserot.

PHOLIDOSIS. Scale rows: 19. Ventrals: 165. Subcaudals: 62.
MEASUREMENTS. SV: 190 mm; tail: 45 mm.

REMARKS. Three juveniles from Birketen near Jerash (HUJ-R 3024, 3063, 3071 ;
November 1945, Coll. Haas and Hoofien) have 164-166 ventrals and 65-67 sub-
caudals. In Cisjordan (N =9), similarly, 160-169 ventrals and 56-66 subcaudals
have been counted.

244 Y. L. WERNER

Coluber rhodorhachis rhodorhachis Jan

(PI. 6 A, B)

Zamenis rhodorhachis Jan, 1865. In De Filippi, Viagg. in Persia, p. 356 (Iran; restricted by

Kramer and Schnurrenberger, 1963, p. 501, to Schiras, Central Persia.)
Coluber rhodorhachis, Parker, 1931. Ann. Mag. Nat. Hist., (10), 8 : 516.

Coluber rhodorhachis rhodorhachis, Khalaf, 1959. Reptiles of Iraq with some notes on the
Amphibians, Baghdad, pp. 75-76.

MATERIAL EXAMINED (i). BM 1965.805 $ Petra, 1965, D. Western.
PHOLIDOSIS. Scale rows: 19. Ventrals: 242. Subcaudals: 133.
MEASUREMENTS. SV: 790 mm; tail: 320 mm.

COLORATION. Dark crossbands on anterior part of back (65-70) somewhat
irregular, nearly four times as broad as the light intervening spaces. First dark
crossband (on the occiput) interrupted mid-dorsally by a faint light vertebral line
(PI. 6 A).

REMARKS. This is apparently the first formal record of C. rhodorhachis from
Transjordan proper. The range extension involved is only minor, as the species is
well known in the Wadi 'Arava (Haas, 1951).

Although this specimen was collected at Petra, and its pholidosis agrees with
material from the adjacent territory of Cisjordan, its coloration deviates markedly
from that found in these specimens. In Cisjordan the dark crossbands are usually
distinctly narrower than the intervening light spaces (PI. 6 B), or at the most as
broad as the latter. The same appears to be true of Egyptian specimens (Anderson,
1898: pi. 35). On the other hand, I have seen broad and close dark crossbands, like
those of the Petra specimen, in an example from Iran (MCZ 58872), though in this
case only the anterior 30-35 crossbands are so broad, the more posterior ones
gradually become narrower. The similarity of these two specimens does not,
however, mean very much, as the species exhibits high variability of colour and
pattern.

Terent'ev and Chernov (1949 : 242) accept C. r. ladacensis Anderson 1871
(Boulenger, 1890 : 326) as a distinct form. Mertens (1956 : 95) and Kramer and
Schnurrenberger (1963 : 501) doubt its validity. In fact, Anderson himself (1895 :
654, footnote i) says "I am indebted to the Trustees of the Indian Museum for the
opportunity of re-examining the types of Z. ladacensis. They are unquestionably
identical with Jan's Z. rhodorhachis. At the time I described the species, Jan's
work was not in the library of the Indian Museum, Calcutta". Thus, so far all
Coluber rhodorhachis specimens, apart from the Somalian C. r. subnigra Boettger
1893 (Parker, 1949 : 30-37), are assignable to the typical form.

Coluber rogersi Anderson
(PI. 6 C, D)

Zamenis rogersi Anderson, 1893. Ann. Mag. Nat. Hist. (6)12 : 439 (Desert to the east of

Helwan, near Cairo).
Coluber rogersi, Flower, 1933. Proc. Zool. Soc. London, p. 810-811.

LIZARDS AND SNAKES FROM TRANSJORDAN 245

MATERIAL EXAMINED (2). BM 1965.698 $ Five km S of Aseikhim, April-May
1965, S. Bisserot. BM 1965.806 juv. $ (?) Azraq, 1965, D. Western.

PHOLIDOSIS. Scale rows: 19; 19. Ventrals: 195; 204. Anals divided. Tails
incomplete.

MEASUREMENTS. SV: 565; 220 mm. (Tails incomplete.)

COLORATION. In both specimens, the anterior three-quarters of body has about
55 dark, closely set, dorsal blotches (PI. 6 C). Posteriorly the blotches gradually
become indistinct. The first three blotches are confluent mid-dorsally to form a
longitudinal streak behind the occiput. Tail uniformly grey.

REMARKS. The pattern resembles that normally encountered in (southern)
Cisjordan (PI. 6 D), the blotches being closer to each other than those figured by
Anderson (1898: pi. 36) for a male from "Beltim" (Nile delta). The scale count
appears to be higher in Transjordan than in Cisjordan: Two specimens from 65 km
SSE of Amman have 200 ventrals each (HUJ-R 3183/1 and 2) ; one from N of Zerka
has 195. Thus while in Cisjordan we find 188-200 ventrals (N = 13), we find
195-204 (N = 5) in Transjordan. A single specimen from Iraq has 206 (HUJ-R
3540).

Malpolon moilensis Reuss

Coluber moilensis Reuss, 1834. Mus. Senckenb. i, p. 142 (near Moila, on the Red Sea coast of

Arabia) .
Malpolon moilensis, Parker, 1931. Ann. Mag. Nat. Hist. (10) 8 : 522.

MATERIAL EXAMINED (i). BM 1965.697 <$ (?). Three miles N of Druze (Track
through hamada), April-May 1965, S. Bisserot.

PHOLIDOSIS. Scale rows: 17. Ventrals: 166. Anal divided. Subcaudals: 50.
MEASUREMENTS. SV: 443 mm; tail: 95 mm.

COLORATION (After preservation). Brown with darker brown spots, of varying
distinctness, which are arranged in eight longitudinal rows. Neighbouring spots
tend to merge, forming obliquely transverse streaks. A conspicuous dark brown
blotch is present on each temporal region.

REMARKS. This appears to be the first record of this snake from Transjordan.
The species has recently been recorded from southern Cisjordan, where it is rare
(Barash & Hoofien, 1956). Most, if not all, specimens were actually found in the
Wadi 'Arava, so that the inclusion of this species in the fauna of Transjordan would
have been a matter of course. The locality recorded here, however, constitutes a
significant range extension.

Malpolon monspessulanus insignitus Geoffroy

Coluber insignitus Geoflroy in Savignyi, 1827. Descr. Egypte., Hist, nat., i Rept. : 151; pi. 7,

Fig. 6 (Lower Egypt).
Malpolon monspessulanus insignitus, Mertens and Miiller, 1928, Abh. senckenberg. naturf. Ges.,

Frankfurt a.M., 41 : 51.

246 Y. L. WERNER

MATERIAL EXAMINED (i). BM 1965.807 <$ Azraq, 1965, D. Western.
PHOLIDOSIS. Scale rows: 19. Ventrals: 173. Anal divided. Subcaudals: 83.
MEASUREMENTS. SV: 745 mm. Tail: 210 mm.

COLORATION (After preservation). Nearly uniform dark grey. Belly whitish
with some light grey mottling.

DISCUSSION

The material reported here does not on its own permit an analysis of the herpeto-
fauna of Trans Jordan in terms of ecology or zoogeography. However, the specimens
have been assembled on several trips, so that at least the lizard material probably
reflects to some extent the abundance of the species in certain habitats and localities.
It thus seems profitable to relate the data available to existing general information
on the ecological and biogeographical subdivision of Trans Jordan and to the known
circumstances in Cisjordan.

General Biogeography of Trans Jordan

The variegated zoogeography of Transjordan was outlined by Bodenheimer
(1935 : 24). His basically correct map (1935: Fig. 6) was superseded by the more
recent work of botanists (Bodenheimer, 1953 : 85). Feinbrun and Zohary prepared
a phytogeographical map (1955 : 15) dividing Transjordan into the same three
territories, Mediterranean, Irano-Turanian, and Saharo-Sindian, into which Cis-
jordan is also divided (Zohary, 1955). This division, based on the mapping of plant
associations (Feinbrun and Zohary, 1955 : folding map), is related to the distribution
of soil types and, more closely, to that of rainfall (maps, Feinbrun and Zohary, 1955 :
9 and 13; Poore and Robertson, 1964 : 12; Bender, 1968 : 10 and 180). The phyto-
geographical map of Feinbrun and Zohary (1955 : 15) furnished the basis for the
delimitation of biogeographical territories in Text-fig. 8. (A fourth territory, the
'Sudanian Penetration Territory', has been proposed for the Lower Jordan Valley
and the Wadi 'Arava by Gruenberg-Fertig, 1965). The general ranges of the three
biogeographical territories in southwestern Asia are presented by a map recently
published elsewhere (Haas and Werner, 1969 : 368).

Poore and Robertson (1964 : 14-15) similarly classified the Transjordanian range
types into three basic regions : Mediterranean, Steppe, and Desert of varying type
(limestone, basalt, sandstone and granite deserts). The salient differences between
the two maps are few : The Mediterranean region of Poore and Robertson is narrower
(in W-E direction) than that of Feinbrun and Zohary (shown here in Text-fig. 8) so
that for instance, Petra, Tafila and Shaubak are excluded from it (the first lying
just west of, the two last, just east of, Poore and Robertson's Mediterranean region).
The Steppe region of Poore and Robertson, roughly coinciding in the south with
Feinbrun and Zohary's Irano-Turanian territory, reaches in the north eastwards to
only halfway between Amman and Azraq. (Guest (1966: Figs 13, 14) apparently
includes the area around, and east of, Azraq, in the Irano-Turanian, in agreement

LIZARDS AND SNAKES FROM TRANSJORDAN

247

FIG. 8. The biogeography of Transjordan, based on Feinbrun and Zohary (1955). The
100 mm mean annual precipitation line is based on Poore and Robertson (1963) ; its inter-
rupted part (extrapolated alternatives in area lacking data) is based on various sources.
M, Mediterranean; IT, Irano-Turanian ; SS, Saharo-Sindian.

248 Y. L. WERNER

with Feinbrun and Zohary.) At the latitude of the Dead Sea Poore and Robertson's
Steppe region has a south-eastern extension (as compared to Feinbrun and Zohary's
Irano-Turanian territory) so as to include El Qatrane. The desert regions of Poore
and Robertson include areas excluded from Feinbrun and Zohary's Saharo-Sindian
territory. Firstly, the district around, and east of, Azraq ; secondly, a broader zone
in the Jordan Valley and especially in its northern part. Thus, Salt (NW of Amman)
and Petra are well within the Mediterranean territory of Feinbrun and Zohary, but
on the fringe of the desert according to Poore and Robertson.

The bioclimates of Transjordan have been defined and mapped by Long (1957;
also in Poore and Robertson, 1964 : 10, and 13). His map corresponds closely to the
two biogeographical maps just discussed; concerning the district surrounding, and
east of, Azraq, it is intermediate between them. The climates of the Mediterranean
territory are "sub-humid and semi-arid mediterranean bioclimates"; those of the
Irano-Turanian (~ Steppe) territory are "arid mediterranean bioclimates"; and the
Saharo-Sindian (~ desert) territory has "saharan mediterranean bioclimates".
Among the latter, the "cool variety" characterizes the Azraq district. Long's map
differs from the phytogeographical maps chiefly in that his "semi-arid mediterranean
bioclimate" (~ Mediterranean territory) reaches southwards only to a point between
Shaubak and Petra. However, this difference may conceivably result from the
paucity of meteorological information available to Long (Poore and Robertson,
1964 : n).

Thus Transjordan comprises three gross ecological, hence biogeographical terri-
tories. Relying on the sources cited, these may be briefly characterized as follows :

The Mediterranean territory includes the hills of Gilead and the western parts of
Ammon, Moab and N Edom. In the south this narrow territory is confined to the
higher hills, and interrupted between them. Predominant soils are terra-rossa,
white Cenomanian soil and grey Senonian soil; along the Jordan Valley, areas of
Nubian sandstone are included. The bioclimate conforms in the main to the semi-
arid mediterranean type. Annual rainfall varies from about 700 mm on northern
mountaintops down to ca. 300, and even below 200, south of Petra. Vegetation is
characterized by a climax of Mediterranean Maquis and forest types, and by areas
covered with various shrubs, including many Labiatae.

The Irano-Turanian territory surrounds the Mediterranean territory except in
the north (where the latter continues into Syria and Lebanon). Above latitude
31 30' N the otherwise narrow zone widens eastwards, extending to at least halfway
between Amman and Azraq, possibly to the frontier. The commonest soils are loess
and grey calcareous steppe soils (Basalt from Azraq to Burqu). The bioclimate is of
the arid mediterranean types (of the saharan mediterranean type, cool variety,
around Azraq) ; annual rainfall is between ca. 350 mm and ca. 100 mm. Vegetation
is typically a steppe of dwarf -shrubs (Artemisia herba-alba is prominent) or of herbs
(e.g. Poa sinica and Car ex pachystylis) , and includes remnants of a Pistacia atlantica
forest.

The Saharo-Sindian territory occupies the vast southern and eastern desert of
Transjordan, as well as the lower parts of the Rift Valley. The greatest part, north
of latitude 29 40' N, is characterized by lime-stone hamadas, the ground generally

LIZARDS AND SNAKES FROM TRANSJORDAN 249

being covered by a layer of flints. Further south, and in the Rift Valley, Nubian
sandstone and sand predominate, with an area of granite (rock and sand) in the Wadi
'Arava. A vast basalt desert, from Azraq to Burqu, is largely covered by basalt
boulders of varying size. The bioclimates are of the Saharan Mediterranean types.
Rainfall is normally below 150 mm, in the extreme southeast below 50 mm. Vegeta-
tion is scanty except in the wadis. Associations of Anabasis articulata are promi-
nent, and on granite sands Haloxylon spp.

Distribution of Reptiles

This discussion is limited to the 23 species and subspecies reported here. Among
these, 16 occur also in Cisjordan. Two ubiquitous species were collected in all three
regions of Transjordan (and similarly occur in all parts of Cisjordan) : Hemidactylus
turcicus and Chalcides o. ocellatus. These have an essentially circum-mediterranean
distribution, which is particularly broad to the south and east. Agama stellio is
similarly circum-eastern-mediterranean, but unfortunately its infraspecific taxonomy
is not clear.

Four forms occur in Cisjordan as Mediterranean elements: Chamaeleo chamaeleon
recticrista, Mabuya vittata, Natrix t. tessellata and Malpolon m. insignitus. The first
three are here reported from the Mediterranean territory of Transjordan; Mabuya
vittata and Natrix t. tessellata were also found at Azraq or at the neighbouring
Shishan (within the disputed area belonging to either the Irano-Turanian or Saharo-
Sindian territory). Malpolon m. insignitus is reported from Azraq. This distribu-
tion is probably due to the local conditions prevailing at these places. N. t. tessellata
is semiaquatic; M. vittata is facultatively hydrophilic, and lives, e.g., also among
reeds surrounding the salt marshes south of the Dead Sea (pers. obs.).

Seven reptiles typical of Cisjordanian desert habitats were collected in the Irano-
Turanian and Saharo-Sindian areas of Transjordan: Agama sinaita, Acanthodactylus
b. asper, Eremias g. gutulata, Eumeces s. schneideri, Coluber rhodorhachis, C. rogersi,
and Malpolon moilensis. We shall consider first the two Coluber species. All of 4
localities for Coluber rogersi fall within the Irano-Turanian (according to Feinbrun
and Zohary; but 2 within the desert according to Poore and Robertson.). In Cis-
jordan this snake occupies both the Irano-Turanian and Saharo-Sindian territories,
with the exception of the (hot) Wadi 'Arava. Its world distribution is Saharo-
Sindian and to some extent, Irano-Turanian. C. rhodorhachis occurs in Cisjordan
mainly in the Saharo-Sindian territory (Wadi 'Arava and southern Negev), and the
first and only record for Transjordan is from Petra, on the fringe of the Wadi 'Arava.
Its world distribution is Irano-Turanian and Saharo-Sindian. The two lacertids,
A. b. asper and E. g. guttulata, are reported from both Irano-Turanian and Saharo-
Sindian localities in Transjordan. Similarly, they are known in Cisjordan from
suitable habitats throughout the Negev. Their world distribution is Saharo-
Sindian. The remaining forms, Agama sinaita, Eumeces s. schneideri and Malpolon
moilensis, were taken in Transjordan at more or less desertic localities (at least
according to Poore and Robertson). In Cisjordan these are restricted to the
Saharo-Sindian part of the Negev, and their world distribution is Saharo-Sindian.

250 Y. L. WERNER

The ranges of the two forms of Ptyodactylus apparently fail to coincide with any of
the ecological territories described above. P. h. puiseuxi, a morphologically well
denned form, is common in northernmost Cis Jordan and in Trans Jordan, on the hills
bordering Lake Tiberias on the east, and around Jerash all in the mediterranean
territory but is here reported also from the fringe of the basalt desert ca. lokm E
of Azraq (Wadi Aseikhim). It may eventually turn out to be basically a form
inhabiting basalt rocks, which has spread to adjacent habitats. P. h. guttatus is less
well defined and in particular not clearly distinguishable from P. h. hasselquistii.
Geckos currently assigned to guttatus are common in the Mediterranean, Irano-
Turanian and Saharo-Sindian territories of Cisjordan in effect throughout the
country excepting its northern and southern extremes. So far, our records for
Transjordan are all within the (southwestern) Saharo-Sindian (Rum; S of Guweira)
or on its rim (Petra; Wadi Musa). Evidently the ranges of these two geckos are
influenced to a great extent by factors other than the climate, perhaps because they
can modify their exposure to it by varying the relative duration of nocturnal and
diurnal activity.

Of the Transjordanian reptiles reported here, seven forms do not occur in Cisjordan.
Of these, six are apparently Irano-Turanian elements, as far as their general ranges
are concerned: Agama p. haasi, Acanthodactylus grandis, A. t. tristrami, Eremias b.
microlepis, Ophisops e. blanfordi, and Eumeces s. princeps. In Transjordan, A. p.
haasi has been collected mostly within the Irano-Turanian, but also within the
Saharo-Sindian. In Cisjordan, A. p. pallida similarly occurs in both territories.
A. grandis has been collected in the Irano-Turanian and Saharo-Sindian, and A. t.
tristrami in the Irano-Turanian and Mediterranean. Some of the localities for E. b.
microlepis are within the Irano-Turanian, the rest within the Saharo-Sindian.
These three lacertids have no conspecific relations in Cisjordan. 0. e. blanfordi has
been collected within the Irano-Turanian and on both its mesic and desertic borders.
The related 0. e. ehrenbergi occurs in the Mediterranean and Irano-Turanian of
Cisjordan, and also in the Mediterranean of Transjordan (around Jerash).

The case of ". s. princeps is of particular interest since three distinct, apparently
allopatric, races of E. schneideri occur in Transjordan and adjacent areas (Mertens,
1920, 1924, 1946; Taylor, 1935 ; Eiselt, 1940). Both localities reported here for E. s.
princeps are within the Irano-Turanian, and this agrees with this race's general
distribution. In the Saharo-Sindian of both Cis- and Transjordan E. s. schneideri
occurs, while E. s. pavimentatus lives in the Mediterranean of Cisjordan. Its
occurrence in Transjordan, which is probable, remains to be shown.

The last of the Transjordanian reptiles not occurring in Cisjordan is Agama
blanfordi fieldi (A. persica fieldi Haas and Werner, 1969). This obviously is a
Saharo-Sindian, Arabian, form. Its taxonomic relationship with the superficially
similar psammophile, A. savignii of eastern Egypt, Sinai and southern Cisjordan,
has not been studied but the two probably occupy comparable ecological niches.

In conclusion, the locality data presented here for Transjordanian lizards and
snakes are in good agreement with a generalized subdivision of Transjordan into
three major ecological-biogeographical territories, based on both the maps of Fein-
brun and Zohary (1955 : 15) and Poore and Robertson (1964 : 14-15). The species

LIZARDS AND SNAKES FROM TRANSJORDAN 251

of the Mediterranean territory exhibit no marked morphological deviations from
their conspecific counterparts in the Mediterranean of Cisjordan. On the other hand,
in several species of desert reptiles (Saharo-Sindian or Saharo-Sindian and Irano-
Turanian), the Transjordan population appears to differ from the Cisjordanian one.
Furthermore, one Saharo-Sindian species, Agama Uanfordi, does not occur west of
the Rift Valley. Conspicuous is the occurrence in Transjordan of six Irano-
Turanian species, of which three do not occur in Cisjordan, while the remaining three
are represented there by other (well defined) subspecies (see also Haas, 1952). The
reciprocal phenomenon also exists as not all Cisjordanian reptiles occur in Trans-
Jordan. Thus among the Saharan psammophile reptiles of southern Cisjordan, five
occur only west of the Rift Valley. Another species, Sphenops sepsoides, penetrates
into the Rift Valley (Werner, 1968), and only one other, Acanthodactylus scutellatus
scutellatus, is represented east of the Rift Valley by another subspecies, A. s. hardyi
(northwestern Saudi Arabia Haas, 1957; Iraq Haas and Werner, 1969). It is
tempting to assume that the Wadi 'Arava, together with the very steep mountain
slopes bordering it on the east, constitutes a barrier to the distribution of reptiles,
though more direct evidence on this effect would be desirable.

ACKNOWLEDGEMENTS

For the privilege of examining this material I am grateful to A. G. C. Grandison,
British Museum (Nat. Hist.). I am much obliged to S. Bisserot who kindly presented
his field notes for inclusion in this report; to D. Western for his cooperation and
stimulating correspondence; and to K. Klemmer, Senckenberg Museum, who most
helpfully supplied information on the type of Agama pallida Reuss, photographs of
it, and a copy of its description.

I wish to thank S. Furman and N. Shulman for assistance in examining specimens;
E. Alcalay and T. Sheffer for help in preparing the diagrams and maps; E. Haupt
for the photographs of Agama pallida Reuss (type) ; E. Ben-Hur and A. B. Niv for
the other photographs; and my wife, Nurit, for both advising me on statistics and
performing the calculations.

I am indebted to E. N. Arnold, British Museum (Nat. Hist.), G. Haas and,
particularly, to J. H. Hoofien, for their painstaking and instructive comments on
the text and illustrations, most of which I heeded.

APPENDIX

LIST OF LIZARDS AND SNAKES SO FAR RECORDED FROM
TRANSJORDAN (INCL. WADI 'ARAVA)

The documentation cited in parentheses, for species not represented in this report,
is not necessarily the earliest one available. The names listed are not necessarily
those employed for the same taxa by the authors cited.

252 Y. L. WERNER

LACERTILIA
GEKKONIDAE

1. Bunopus blanfordii Strauch (Barash and Hoofien, 1956)

2. Ceramodactylus doriae Blanford (Haas, 1956)

3. Hemidactylus turcicus turcicus L.

4. Pristurus flavipunctatus guweirensis Haas (Haas, 1951)

5. Ptyodactylus hasselquistii guttatus von Heyden

6. Ptyodactylus hasselquistii puiseuxi Boutan

7. Stenodactylus grandiceps Haas (Haas, 1951)

8. Stenodactylus sthenodactylus sthenodactylus Lichtenstein (Haas, 1951)

9. Tropiocolotes steudneri Peters (Haas, 1951)

AGAMIDAE

10. Agama blanfordi fieldi Haas and Werner

11. Agama pallida haasi Werner

12. Agama sinaita von Heyden

13. Agama stellio brachydactyla Haas

14. Agama stellio picea Parker (Haas, 1951)

15. Agama stellio stellio L. (Daan, 1967)

CHAMAELEONIDAE

16. Chamaeleo chamaeleon recticrista Boettger

LACERTIDAE

17. Acanthodactylus boskianus asper Audouin

18. Acanthodactylus cantoris schmidti Haas (Hoofien, 1965)

19. Acanthodactylus grandis Boulenger

20. Acanthodactylus robustus Werner (Haas, 1951)

21. Acanthodactylus tristrami tristrami Giinther

22. Eremias brevirostris microlepis Angel

23. Eremias guttulata guttulata Lichtenstein

24. Eremias guttulata watsonana Stoliczka? (Wettstein, 1951)

25. Eremias olivieri schmidti Haas, (Haas 1951)

26. Lacerta danfordi danfordi Giinther (Hoofien, 1969)

27. Ophisops elegans blanfordi Schmidt

28. Ophisops elegans ehrenbergi Wiegmann (Haas, 1951)

29. Ophisops elegans elegans Menetrie's (Schmidt, 1939)

SCINCIDAE

30. Ablepharus kitaibelii Bibron et Bory (Haas, 1951)

31. Chalcides ocellatus ocellatus Forskal

32. Eumeces schneideri princeps Eichwald

LIZARDS AND SNAKES FROM TRANSJORDAN 253

33. Eumeces schneideri schneideri Daudin

34. Mabuya vittata Olivier

35. Ophiomorus latastii Boulenger (Anderson & Leviton, 1966)

36. Scincus sp. (Haas, 1951)

37. Sphenops sepsoides Audouin (Werner, 1968)

ANGUIDAE

38. Ophisaurus apodus Pallas (Haas, 1951)

VARANIDAE

39. Varanus griseus griseus Daudin (Haas, 1951)

OPHIDIA

LEPTOTYPHLOPIDAE

40. Leptotyphlops phillipsi Barbour (Haas, 1951)

COLUBRIDAE

41. Coluber jugularis jugularis L. (Haas, 1951)

42. Coluber najadum Eichwald (Haas, 1951)

43. Coluber ravergieri nummifer Reuss (Flower, 1933)

44. Coluber rhodorhachis Jan

45. Coluber rogersi Anderson

46. Eirenis collaris Me'ne'trie's (Haas, 1951)

47. Eirenis coronella coronella Schlegel (Schmidt, 1939)

48. Eirenis coronella fraseri Schmidt (Haas, 1951)

49. Eirenis decemlineata Dumeril & Bibron (Haas, 1951)

50. Eirenis rothi Jan (Haas, 1951)

51. Malpolon moilensis Reuss

52. Malpolon monspessulanus insignitus Geoffroy

53. Natrix tessellata tessellata Laurenti

54. Psammophis schokari Forskal (Haas, 1951)

55. Rhynchocalamus melanocephalus Jan (Hart, 1891)

56. Spalerosophis diadema ssp. (Haas, 1951)

57. Tarbophis nigriceps Ahl (Haas, 1951)

VIPERIDAE

58. Cerastes cerastes L. (Haas, 1951)

59. Echis colorata Gunther (Haas, 1951)

60. Pseudocerastes fieldi Schmidt (Haas, 1951)

254 Y. L. WERNER

REFERENCES

ANDERSON, J. 1895. On a collection of reptiles and batrachians made by Colonel Yerbury

at Aden and its neighbourhood. Proc. zool. Soc. Lond. 1895 : 635-664.
- 1896. A contribution to the herpetology of Arabia. With a preliminary list of the reptiles
and batrachians of Egypt. R. H. Porter, London, 122 pp.

1898. Zoology of Egypt. Vol. i, Reptilia and Batrachia. Quaritch, London, LXV +

37i PP-
ANDERSON, S. C. ig66a. A substitute name for Agama persica Blanford. Herpetologica,

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LIZARDS AND SNAKES FROM TRANSJORDAN 255

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256

Y. L. WERNER

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REFERENCES TO APPENDIX

ANDERSON, S. C. & LEVITON, A. E. 1966. A review of the genus Ophiomorus (Sauria:
Scincidae), with descriptions of three new forms. Proc. Calif. Acad. Sci., Ser. 4, 33 (16) :

499-534-
HAAS, G. 1956. On the occurence of Stenodactylus petrii and Stenodactylus (Ceramodactylus)

doriae in southern Israel. Bull. Res. Coun. Israel, 5B (3-4) : 308-309.
HART, H. C. 1891. Some account of the fauna and flora of Sinai, Petra, and Wddi 'Arabah.

Watt, London, X + 255 pp., folding map.

Dr. Y. L. WERNER

Department of Zoology

HEBREW UNIVERSITY OF JERUSALEM

JERUSALEM, ISRAEL

PLATE i

Agama pallida subspp. :

(A-D) A. p. haasi subsp. nov. : (A) Holotype^, BM 1965.800, dorsal view; (B) Same, left ear;
(C) Juvenile, BM 1965.797, right ear; (D) Paratype $, HUJ-R 1884.

(E-F) A. p. pallida Reuss : (E) From southern Cisjordan (Wadi Ajram) <$, HUJ-R 1623;
(F) From eastern Egypt (Kassassin) $, HUJ-R 1126.

Scale, cm and mm (D-F at same magnification).

Bull. Br. Mus. nat. Hist. (Zool.) 21,6

PLATE i

0123

CH & 8 L

PLATE 2

Agama pallida pallida Reuss holotype $, SMF 10007:
(A) Dorsal view; (B) Head.

Bull. Br. Mus. not. Hist. (Zool.) 21, 6

PLATE 2

PLATE 3

(A) Agama blanfordi fieldi $ , BM 1965.686.

(B-E) Agama sinaita: (B) Dorsal view of left pes, and (C) of base of tail, of ^ from northern
Transjordan (Wadi Ratam), BM 1965.685; (D) Dorsal view of left pes of $ from southern
Cisjordan, HUJ-R 1919; (E) Dorsal view of tail base of <$ from southern Cisjordan, HUJ-R 1794.
Scale, cm and mm (B-E at same magnification).

Bull. Br. Mus. nat. Hist. (Zool.) 21,6

PLATE 3

PLATE

(A-B) Acanthodactylus grandis : (A) Adult $ from Tell el Mukheizin, BM 1965.694; (B)
Young $ from Ain el Enoquiyya, BM 1965.692.

(C-D) Acanthodactylus tristrami tristrami : (C) $ from the Jordanian-Syrian border, BM
1962.352; (D) 3 1 from 15 Km S of Amman, HUJ-R 1333.

Scale, cm and mm (all at same magnification).

Bull. Br. Mus. nat. Hist. (Zool.) 21,6

PLATE 4

PLATE 5

Eumeces schneideri subspp. :

(A) E. s. schneideri from Petra, BM 1965.786; (B) E. s. princeps from Ain el Enoquiyya,
BM 1965.695; (C) E. s. schneideri from N Shaubak, BM 1963.669.

Scale, cm and mm.

Bull. Br. Mus. nat. Hist. (Zool.) 21, 6

PLATE 5

PLATE 6

(A-B) Coluber rhodorhachis rhodorhachis: (A) <j> from Petra, BM 1965.805; (B) <$ from southern
Cisjordan, HUJ-R 3211.

(C-D) Coluber rogersi : (C) $ from 5km S of Aseikhim, BM 1965.698; (D) 9 from southern
Cisjordan, HUJ-R 8020.

Scale, cm and mm.

Bull. Br. Mus. nat. Hist. (Zool.) 21,6

PLATE 6

:j I i II j II i 1 1 1 1 1 1 1 i 1 1 1
OARISTO1 2

A LIST OF SUPPLEMENTS
TO THE ZOOLOGICAL SERIES

OF THE BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

1. KAY, E. ALISON. Marine Molluscs in the Cuming Collection British Museum
(Natural History) described by William Harper Pease. Pp. 96; 14 Plates.
1965. (Out of Print.) 3.75.

2. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier and
Valenciennes. Pp. 180; n Plates, 15 Text-figures. 1967. 4.

3. TAYLOR, J. D., KENNEDY, W. J. & HALL, A. The Shell Structure of Mineralogy
at the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125; 29 Plates,
77 Text-figures. 1969. 4.50.

4. HAYNES, J. R. Cardigan Bay recent Foraminifera (Cruises of the R.V. Antur)
1962-1964. (In press.)

Printed in England by Staples Printers Limited at their Kettering, Northants, establishment

i \ CONCHOECIA FROM THE
NORTH ATLANTIC

THE 'PROCERA' GROUP

M. V. ANGEL

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 21 No. 7

LONDON: 1971

CONCHOECIA FROM THE NORTH ATLANTIC
THE 'PROCERA' GROUP

MARTIN VIVIAN ANGEL V|J

/NU

National Institute of Oceanography

Pp. 257-283 ; 14 Text-figures

THE BRITISH MUSEUM (NATURAL HISTORY)

ZOOLOGY Vol. 21 No. 7

LONDON: 1971

THE BULLETIN OF THE BRITISH MUSEUM

(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical series.

Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.

In 1965 a separate supplementary series of longer
papers was instituted, numbered serially for each
Department.

This paper is Vol. 21, No. 7 of the Zoological
series. The abbreviated titles of periodicals cited follow
those of the World List of Scientific Periodicals.

World List abbreviation
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Trustees of the British Museum (Natural History), 1971

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Issued 3 December, 1971 Price 1.05

CONCHOECIA FROM THE NORTH ATLANTIC
THE 'PROCERA' GROUP

By MARTIN VIVIAN ANGEL

INTRODUCTION

MULLER (1894) decided that Conchoecia variabilis Miiller 1890 was a confusion of
two species, C. oblonga Claus 1890 and a new species C. procera. Claus (1894)
also confused these two species. Later Muller (i9o6a) further described C, procera,
and gave its size range as 1-05-1-35 mm for females and 0-85-1-20 mm for males.
The geographical range was described as being between 3iN and 35 S in all oceans
(Muller I9o6a, b, 1908, 1912). It has since been further reported in the North-
eastern Atlantic (Fowler 1909, Granata & Caporiacco 1949, Angel 1968, iQ6ga., b),
the Sargasso Sea (Deevey 1968), the Benguela Current (lies 1953), western Mediter-
ranean (Leveau 1965), the Adriatic (Schweiger 1912, Mure" 1955, 1961), the western
Indian Ocean (Leveau 1967, 1968), the eastern Indian Ocean (McKenzie in press).

In 1968 R.R.S. 'Discovery' took a series of hauls in the region of nN 20W,
about 300 miles south of the Cape Verde Islands, to study vertical distribution
patterns. It was soon apparent that adult specimens attributable to C. procera
separated into three distinct size groupings, which showed differential depth distri-
butions; Deevey (1968) also noted two size groupings. The middle sized group, the
most abundant, is considered here to be synonomous with C. procera sensus strictu.
The morphological differences between this group and the smaller and larger groups
are sufficiently great to attribute them with specific status. They are, therefore,
named Conchoecia microprocera sp. nov. and Conchoecia macroprocera sp. nov.
respectively.

A single adult female taken in a deep vertical haul at 'Discovery' station 4768
in 1961 near 40 N 2OW, is also described and is ascribed to C. vitjazi Rudjakov
1962, another species which possibly belongs to the 'procera' group.

Conchoecia procera Muller 1894

MALE. Carapace. The lengths of 234 specimens from nN 20W ranged from
0-98-1-04 mm with a mean of 1-020 + 0-019 mm, and of 70 specimens from off the
Moroccan coast 34N 8W (Angel 1968) ranged from 0-96-1-04 mm with a mean of
1-005 0-009 mm - The outline of the carapace (fig. lA, B) with the smooth curve
of the posterior edge into the ventral edge broken only by the step at the opening
of the right asymmetrical gland is similar to Miiller's figure (1894 T. XIII fig. 39).
Similarly the shape of the posterior dorsal spine (fig. 3F) is identical to Miiller's
figure (1894 T. XIII fig. 41) and is the character by which he separated C. procera
from C. decipiens Muller in his key (Muller 1912). Edge glands are numerous down
the posterior carapace margin, and are present but less abundantly along the

260

M. V. ANGEL

FIG. i. Conchoecia procera Muller, male. A, Lateral view of carapace; B, Ventral view
of carapace; C, Frontal organ and first antenna; D, Armature of the b and e setae of the
first antenna; E, sixth limb; F, second antenna; G, endopodite of the right second
antenna; H, left hook appendage; I, right hook appendage; J, penis; K, detail of tip
of penis.

CONCHOECIA FROM THE NORTH ATLANTIC 261

ventral margin. Live animals have not been observed to bioluminesce.

Frontal organ (fig. iC). The shaft reaches level with the end of the limb of the
first antenna. The capitulum is downturned and slightly curved. There is a small
ventral swelling about half way along its length.

First antenna (fig. iC). The first segment is shorter than the second. None
of the segments carry any additional armature. The a seta is almost as long as the
limb, reaching well beyond the joint between the first and second segments. The
c seta is very short. The remaining three setae are almost equal with the e seta
just the longest. The d seta is bare but the b seta has three small spines opposite
the distal end of the e seta armature. The e seta has about seventeen pairs of
long slender spines (fig. iD) with two pairs of distally pointing spinules just distal
of the main armature. The main spines decrease in length slightly towards the base
of the seta.

Second antenna (fig. iF). The exopodite segments carry no unusual armature.
The protopodite is a little less than half the carapace length and the first exopodite
segment is about 3/8's its length. On the endopodite (fig. iG) the processus mamil-
laris is blunt with a small rounded tubercle slightly offset to one side. The a seta
which carries fine spinules, is half the length of the b seta. The b seta is armed
with much longer and stronger spines. The c and d setae are almost as long as the
second segment, while the e seta is a minute spine. The f seta is only slightly
shorter than the g seta, and both setae are thin walled terminally and unarmed.
The h, i and j setae are short with barely developed shafts, but the h seta does have
a small basal swelling. The left hook appendage (fig. iH) curves through about
120, terminally tapering into a smooth point. The right hook appendage (fig. iG)
is larger and slightly more curved. It is slightly swollen and ridged subterminally
and ends in a point.

Mandible (fig. 2C). There is only a single long seta on the inner surface of the
first endopodite segment, and there is no additional armature on any of the other
endopodite segments. The toothed edge of the basale (fig. 2B) is typical for the
genus with two tubiform teeth, six serrate teeth and an outer tooth which is unusually
broad in this species. One of the outer setae is long, but the other only reaches
level with the teeth. There are long hairs on the outer surface arranged in four rows,
two leading up to the bases of the spine teeth. The outer teeth have very clear
secondary tooth rows. The toothed edge of the coxale has ten teeth (fig. 2 A).
The distal tooth list consists of two large teeth, neither of which is serrate, and ten
to twelve small teeth, the outermost of which is broad. The proximal list consists
of 13-15 fairly regular teeth which decrease in size along the list.

Maxilla (fig. 2D). There is a basal seta. The first endopodite segment has six
anterior, one lateral and three posterior setae. There is no distal armature on the
first segment.

Labrum (fig. 2E). The hyaline membrane has a smooth rather shallow notch.

Caudal furca (fig. 2G). There is no unpaired seta dorsal to the eight pairs of hook
spines. There is a covering of fine hairs between the caudal lamellae.

Penis (figs ij, K). The intromittent organ has five oblique muscles. The end
of the organ is rounded in outline.

2&2

M. V. ANGEL

FIG. 2. Conchoecia procera Miiller. A, mandible tooth list and toothed edge of the coxale ;
B, mandible toothed edge of the basale; C, mandible endopodite; D, maxilla endopodite;
E, labrum; F, fifth limb; G, caudal furca.

CONCHOECIA FROM THE NORTH ATLANTIC

263

FIG. 3. Conchoecia procera Miiller, female. A, lateral view of carapace; B, ventral view
of carapace; C, frontal organ and first antenna; D, second antenna; E, second antenna
endopodite; F, male detail of the posterior dorsal corner of the carapace.

FEMALE. Carapace (figs 3A, B). The lengths of 282 specimens from nN
ranged from 1-12-1-24 mm with a mean of 1-181 + 0-021 mm. 122 specimens from
off the Moroccan coast ranged from 1-12-1-24 mm with a mean of 1-166 + 0-022 mm.
The change in the mean size with latitude is insignificant.

Frontal organ (fig. 3C). The capitulum is not differentiated from the stalk.
The total length is about three times the length of the limb of the first antenna It
has a slight terminal swelling with a rounded knob. There are spines on its under-
side.

264 M. V. ANGEL

First antenna (fig. 36) . There is no additional armature. The a-d setae are
thin walled and a little longer than the length of the limb. The e seta is nearly
twice as long as the other setae and distal of their ends it carries long fine spinules
on the anterior edge and shorter spinules on its trailing edge.

Second antenna (fig. 3D). The protopodite is 2/5's the length of the carapace
and nearly three times the length of the first exopodite segment. The longest
swimming seta is 3/4 the length of the protopodite. On the endopodite (fig. 3E)
the a seta is bare and less than half the length of the b seta. The b seta carries
8-9 strong spinules. On the segment near the bases of the a and b setae are about
eight small spines. The second segment carries many long fine hairs (c.f. Miiller
1894 T.6 fig. 61). The c, d and e setae are all absent. All the main setae are
thin walled. The g seta is distinctly broader but only slightly longer than the other
four subequal setae.

Synonomy. There is no clear type locality for this species. Miiller (1890)
described C. variabilis which he later (1894) decided was a confusion between C.
oblonga Claus and C. procera, from the Pacific and also from the Gulf of Naples.
Claus's (1894) material described as C. oblonga Claus included C. procera Miiller
(Miiller I9o6a) came from the eastern Mediterranean. It is not until Miiller's
(1894) original description of C. procera that any certainty can be attached to the
identifications. However, there are no localities given for any of the material.
From Leveau's (1965) rather sparse data and some specimens from the North
Adriatic sent to me by Professor J. Stirn it has been possible to confirm the presence
of C. procera sensu strictu in the Mediterranean. Miiller's (igoGa) material included
at least one if not both of the new species described below, judging from his data
on size ranges and the armature of the male antennular e seta. Fowler (1909)
appeared only to have caught C. procera sensu strictu from the Bay of Biscay.
Deevey (1968) reported two forms of C. procera, the larger of which conformed with
the description above of C. procera sensu strictu. A re-examination of Angel's
material from the Moroccan coast (1968) and from near Fuerteventura in the
Canary Islands (i969a) showed that they all beonged to C. procera sensu strictu,
with the single exception of the one adult female caught above the thermocline
at 'Discovery' station 6183 haul 2. This specimen corresponded to the new species
described below C. microprocera.

Conchoecia microprocera sp. nov.

MALE. The holotype mounted on slides in Euparal and stained with lignin
pink has been deposited in the British Museum (Natural History), No. 1971.2.1.1.
Locality 'Discovery' station 6665 haul 4, io32.7'N, i9574'W. Depth 400-
295 m. Time 1559-1731 hrs. Date 22 February 1968. Net Modified Indian
Ocean Standard Net (Nii3) fitted with a catch dividing bucket (Foxton 1963, 1969).

Carapace (figs lA, B). The range in length of 204 specimens was 0-82-0-92 mm
with a mean of 0-863 + 0-013 mm The height and breadth of the carapace were
both approximately 2/5's the length. In C. procera sensu strictu the carapace
height was always measurably larger than its breadth (Table i). There was a small

CONCHOECIA FROM THE NORTH ATLANTIC

265

FIG. 4. Conchoecia microprocera sp. nov., male. A, lateral view of carapace; B, ventral
view of carapace; C, frontal organ and first antenna; D, first antenna armature on e seta;
E, sixth limb; F, second antenna; G, endopodite of the right second antenna; H, left
hook appendage; I, penis.

266 M. V. ANGEL

spine on the right valve at the posterior dorsal corner (fig. 6F). The asymmetrical
glands and the edge glands are very similar in their positions to C. procera sensu
strictu.

Frontal organ (fig. 46). The stalk projects just beyond the length of the first
antenna. The capitulum is long and slender and curves slightly downwards.
Terminally it is rounded. There is a patch of fine long hairs in the centre of the
ventral surface.

First antenna (fig. 46). There is no additional armature. The first and second
segments are subequal, and there are large pigment corpuscles in the region of their
articulation. The a seta lies back parallel with the limb and reaches almost to the
base. It is relatively longer than in the other two species (Table i). The c seta
is extremely short. The b and d setae are subequal, only slightly shorter than the
e seta. Both carry very fine spinules on a level with the distal end of the e seta
armature. The e seta armature (fig. 4D) consists of 11-13 pairs of spines which
increase very slightly in length towards the base of the seta. Beyond these main
spines are two pairs of distally pointing spinules. Deevey (1968) described some of
her specimens of C. procera as having this type of armature.

Second antenna (fig. 4?). The protopodite is more than half the length of the
carapace, and this distinguishes this species from the others (Table i). The first
exopodite segment is a third the length of the protopodite and twice the lengths
of the remaining exopodite segments. The longest swimming seta is about 4/5*5
the length of the protopodite. There is no additional armature on the exopodite
segments. On the endopodite (fig. 40) the processus mamillaris has a very bluntly
pointed tip. The a seta is bare and half the length of the b seta. The b seta carries
spinules for most of its length. The c and d setae are almost as long as the second
segment. The e seta is a minute spine. The g and f setae are bare, terminally
thin walled, blunt and slightly flattened. The h, i and j setae are short with very
poorly developed shafts. The' left hook appendage (fig. 46) is right angled at its
base, and then distally straight. It broadens near its end and then tapers to end
in a smooth curved point. The right hook appendage (fig. 4H) is longer beyond the
basal angle, and curves to terminate in a curved point with marked subterminal
ridging.

Mandible (fig. 56). The toothed edge of the basale is the usual structure for the
genus (fig. 56). The outer two setae are very short not reaching level with the
teeth. There are fine hairs on the outer surface, with two rows running up to the
bases of the tubiform teeth. There are no hairs on the basale between the toothed
edge and the articulation with the first endopodite segment. On the first endo-
podite segment there are two setae on the inner edge; one very long, the other
minute. The terminal claw seta is particularly long.

The toothed edge of the coxale (fig. 5A) has ten teeth. The distal tooth list con-
sists of two large teeth, neither serrate, followed by 13-14 small regularly sized
teeth. The proximal list consists of 15 irregular teeth.

Maxilla (fig. 5D). There is a basal seta. There are three posterior setae and a
lateral seta on the first segment, but there are only five anterior setae compared

CONCHOECIA FROM THE NORTH ATLANTIC

267

FIG. 5. Conchoecia microprocera sp. nov. A, mandible tooth lists and toothed edge of
the coxale; B, mandible toothed list of the basale; C, mandible endopodite; D, maxilla
endopodite; E, labrum; F, fifth limb; G, caudal furca.

268 M. V. ANGEL

with six in the other species of the 'procera' group. There is a row of fine spinules
on the distal edge of the first endopodite segment.

Labrum. There is a shallow smooth notch in the hyaline membrane.

Penis (fig. 4!). There are four oblique muscles. The terminal incurved end plate
has two small spines.

Caudal furca (fig. 5G). There is no unpaired seta dorsal of the hook spines.
There are fine hairs between the two caudal lamellae which are most clearly seen
between the bases of the first and second hook spines.

FEMALE. The paratype specimen mounted on slides in Euparal and stained
with lignin pink has been deposited in the British Museum (Natural History) No.
1971.2.1.2 The collection data is as for the male.

Carapace (figs 6A, B). The range in length of 260 specimens was 0-92-1-06 mm
with a mean of 0-999 0*023 mm - The general appearance was more hyaline
than for the other closely related species. It is slimmer in appearance, but the
tendency for the animals to splay open on preservation probably contributed to the
insignificance in the difference between its relative breadth and those of the other
species. The openings of the various glands were similar to those of the other
species. The spine on the right valve at the posterior dorsal corner is well de-
veloped.

Frontal organ (fig. 6C). The frontal organ shows some evidence of differentiation
into stalk and capitulum. The capitulum is a little broader and is swollen towards
its tip finally ending in a down curved point.

First antenna (fig. 6C). There is no dorsal seta or other additional armature.
The a-d setae are only half the length of the e seta. Distal of their ends the e seta
carries short spinules on its posterior edge and sparse long hairs on its anterior
edge.

Second antenna (fig. 6D) . The protopodite is much longer relative to the carapace
than in the other species (Table i). There is no unusual armature on the exopodite.
On the endopodite (fig. 6E) the processus mamillaris is rounded. The a seta is
half the length of the b seta and carries very fine spinules. The b seta has four
to six stouter spinules, and it curves into a long tapering point. On the segment
near the bases of these two setae are four spines which are much longer than in the
other two species. The c, d and e setae are absent. The main setae are all thin
walled and bare. The g seta is only a little longer than the others but appreciably
broader.

Synonomies. It is clear from Deevey's (1968) account that her material included
specimens of this species; both from the size ranges and the number of spines in
the armature of the e seta of the male antennule of her specimens described as C.
procera Miiller. No other authors give sufficient information to determine the full
range of this species. Miiller (i9o6a) also confused this species with C. procera. A
re-examination of material from the Moroccan coast (Angel 1968) showed that the
single adult female caught above the thermocline at 'Discovery' station 6183 haul 2
(34i4'5'N, o803-o'W) belonged to this species. Recently more specimens were
taken in the vicinity of I745'N, 2530'W (Angel unpublished).

CONCHOECIA FROM THE NORTH ATLANTIC

269

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270

M. V. ANGEL

FIG. 6. Conchoecia microprocera sp. nov., female. A, lateral view of carapace, B, ventral
view of carapace; C, frontal organ and first antenna; D, second antenna; E, second
antenna endopodite; F, male detail of the posterior dorsal corner of the carapace.

Conchoecia tnacroprocera sp. nov.

MALE. The holotype specimen mounted on slides in Euparal and stained with
lignin pink has been deposited in the British Museum (Natural History) No. 1971.2.1.3.
Collection data as for C. microprocera sp. nov. (see above).

CONCHOECIA FROM THE NORTH ATLANTIC

FIG. 7. Conchoecia macroprocera sp. nov., male. A, lateral view of carapace; B, ventral
view of carapace; C, frontal organ and first antenna; D, first antenna detail of armature
of the e, d and b setae; E, frontal organ capitulum; F, second antenna; G, left second
antenna endopodite; H, right hook appendage; I, left hook appendage; J, sixth limb;
K, penis.

272 M. V. ANGEL

Carapace (fig. 7 A). The range in size of 207 specimens was i- 10-1-22 mm with
a mean of 1-153 0-021 mm. The relative height tended to be less and the breadth
greater than for the other species. However, a distinguishing feature is that in all
specimens the carapace breadth was greater than the height. The openings of the
various glands are as described in the other species. There is no spine on the right
valve at the posterior dorsal corner (fig. gF) (c.f. Miiller igo6a T. XIII fig. 37).

Frontal organ (fig. 76). The shaft reaches level with the end of the limb of the
first antenna. The capitulum is long and down turned. Two lateral thickening
bars carry fine long hairs for the proximal third. The ventral edge has a small
swelling on its median third, on the distal end of which is a more obvious group of
long hairs. The dorsal edge is concave and bare. The end of the capitulum is
rounded with a low subterminal knob (fig. 7E). One of Miiller's figures (igo6a
T. XIII fig. 45) shows some resemblance to the frontal organ of this species.

First antenna (fig. 76). The first segment is much shorter than the second;
contrasting with the other two species. The ganglionic body, which lies in the region
of the articulation between the two segments contains many yellow brown pigment
corpuscles. The a seta lies back parallel with the limb reaching beyond the end
of the second segment. The seta is significantly shorter than in C. microprocera,
but not significantly so than in C. procera Miiller. The b and d setae are sub-equal
and only carry six and three minute spinules respectively on a level with the distal
end of the e seta armature. The e seta armature (fig. 7D) consists of 28-30 pairs
of long slightly curved spines which increase in length slightly towards the base of the
seta. Distal of the main armature are two pairs of distally pointing spinules.

Second antenna (fig. 7F). The relative lengths of the protopodite and exopodite
are similar to those in C. procera Miiller. The longest swimming seta is longer and
more closely approaches the protopodite length than in the other species. The
a seta on the endopodite (fig. 7G) is 2/3's the length of the b seta and has a few very
fine spinules. The b seta carries more longer spinules. The c and d setae are nearly
as long as the second segment. The e seta is minute. The g seta is only a little
longer than the f seta. Both the g and f setae are bare and the g seta is slightly
flattened. The h seta has a slight swelling near its base. The left hook appendage
(fig. 7!) has a basal right angle and then curves gently a further 30 or so. Ter-
minally it tapers asymmetrically to a blunt point. The right hook appendage
(fig. 7H) curves through a total of 150. The end is swollen with subterminal
ridging and ending in a blunt asymmetrical point.

Mandible (fig. 8C). The toothed edge of the coxale (fig. 8 A) has ten low blunt
teeth. The distal tooth list consists of two large teeth, the second of which is
serrated, followed by about twelve small regularly sized teeth. The proximal list
has twelve to fifteen teeth regularly diminishing in size across the list.

The toothed edge of the basale (fig. 8B) consists of the usual two tubiform fol-
lowed by six wedge teeth with a single broad outer tooth. The outer setae are short
and fail to reach the level of the teeth. There are five rows of hairs arranged on the
surface between the insertion of the outer setae and the two tubiform teeth. On
the inner edge of the first endopodite segment is a single long seta. The second
segment is bare on its outer edge.

CONCHOECIA FROM THE NORTH ATLANTIC

FIG. 8. Conchoecia macroprocera sp. nov. A, mandible tooth lists and toothed edge of
the coxale ; B, mandible toothed edge of the basale ; C, mandible endopodite ; D, maxilla
endopodite; E, labrum; F, fifth limb; G, caudal furca.

274 M. V. ANGEL

Maxilla (fig. 8D). There is a basal seta. On the endopodite the first segment
carries six anterior, one lateral and three posterior setae. There are about six
spinules on the distal edge of the segment near its articulation with the second
segment.

Labrum (fig. 8E). The hyaline membrane has a shallow rounded notch.

Caudal f urea (fig. 8G). The first hook spines reach just short of the tips of the
second pair. There is no unpaired seta dorsal of the hook spines. Between the
two caudal lamellae is a covering of fine hairs.

Penis (fig. 7K). The end of the organ is rounded and it contains four oblique
muscles.

FEMALE. The paratype mounted on slides in Euparal and stained with lignin
pink has been deposited in the British Museum (Natural History) No. 1971.2.1.4.
Collection data as for C. microprocera sp. nov.

Carapace (figs gA, B). The range in length of 263 specimens was 1-26-1 -36 mm
with a mean of 1-304 + 0-021 mm. The relative height of the carapace is noticeably
less than for C. procera Miiller although its breadth is similar. The asymmetrical
glands are positioned as in the other two species. There is no spine on the right
valve at the posterior dorsal corner.

Frontal organ (fig. gC). The total length of the organ is nearly three times the
length of the limb of the first antenna. There is some demarcation into shaft
and capitulum. Near the rounded tip there is a row of spines on the ventral surface.

First antenna (fig. gC). The segmentation is indistinct and there is no additional
armature. The a-d setae are more than half the length of the e seta. The e seta
carries short spinules on its posterior edge and sparser long spinules on the distal
half of its anterior edge.

Second antenna (fig. gD). The ratio of the relative lengths of the segments of the
limb are very similar to those for C. procera Miiller (Table i), although the longest
swimming seta does tend to be longer. On the endopodite the a seta is bare and
2/3's the length of the b seta. The b seta carries a few short spinules. On the
segment near the bases of these setae are about eight short fine spinules. The
second segment is almost bare with only about three or four hairs. All the main
setae are bare and thin walled. The g seta is only a little longer than the others
but is much broader.

Synonomies. Miiller (igoSa) gave a size range for C. procera which suggests
that his material included this species. Similarly Leveau's (1965) report of C.
procera Miiller reaching a length of 1-30 mm may indicate its presence in the Mediter-
ranean. Leveau reported two centres of abundance in C. procera Miiller at 200-
100 m and 500-300 m. It seems possible that the deeper population may be this
species. The species was absent from Deevey's (1968) material, but has turned up
in very small numbers recently in samples taken near i745'N, 253o'W (Angel
unpublished). Thus this species may not normally occur north of about i8N in
the North Atlantic.

CONCHOECIA FROM THE NORTH ATLANTIC

275

FIG. 9. Conchoecia macroprocera sp. nov., female. A, lateral view of carapace; B, ventral
view of carapace; C, frontal organ and first antenna; D, second antenna; E, second
antenna endopodite; F, details of the posterior dorsal corner of the male carapace.

Conchoecia vitjazi Rudjakov 1962

MATERIAL. A single female was found in a 'Discovery' net (Currie & Foxton
1957) haul from 'Discovery' station 4768, position 4O03'N, I957'W, depth 4750-
4000 m, time 1402-1640 hrs, date 12 October 1961. Since no description is available
in English, the specimen is described in full.

276 M. V. ANGEL

MALE. The male is unknown.

FEMALE. Carapace. The length of the 'Discovery' specimen is 2-00 mm.
Rudjakov's specimens ranged from 2-20-2-30 mm. The 'Discovery' specimen was
too distorted to make accurate measurements of carapace height and breadth,
but the height is about half the carapace length. The whole carapace is covered
with a fine V-shaped sculpturing (fig. loA). The right asymmetrical gland opens
on a small prominence (possibly an artefact) and the left asymmetrical gland about
o-i mm from the posterior dorsal corner (fig. loA). The right valve carries a minute
spine at the posterior dorsal corner. The outline of the carapace is very similar
to Rudjakov's (1962) figure 2a.

Frontal organ (fig. loB). The organ is much longer than the limb of the first
antenna. There is no clear distinction into stalk and capitulum. Terminally the
organ is down-turned and carries a few small spinules on the ventral corner (fig. loC).

First antenna (fig. loB). The segmentation is indistinct and there is no addi-
tional armature. The dorsal seta is absent. The e seta is over twice the length
of the other setae and only carries a few fine spinules on the distal third of its trailing
edge. These spinules were on the third quarter of the e seta in Rudjakov's speci-
mens.

Second antenna (fig. loE). The proportion of the protopodite : exopodite seg-
ment i ; exopodite segments 2-8 is 6 : 3 : i in the 'Discovery' specimen and 4:2-8:1
in Rudjakov's specimens. On the endopodite (fig. loF) the processus mamillaris
is small and bluntly pointed. The a seta is bare and two thirds the length of the
b seta. The b seta carries a group of long hairs near its base and very fine spinules
for about half its length. The c, d and e setae are all absent. The g seta is as long
as the protopodite, and is flattened terminally with fine spinules along the edge
of the flattened part. The other setae are long subequal, thin-walled and without
shafts.

Mandible (fig. nB). The toothed edge of the basale is typical for the genus
(fig. 1 1 A). The broad outer tooth is as broad as three of the other teeth. The
outer setae project just beyond the level of the teeth. There are four groups of
hairs near the bases of the tubiform teeth. Between the toothed edge and the
articulation with the first endopodite segment is a patch of fine hairs. The inner
edge of the first endopodite segment carries one long seta reaching well beyond the
tips of the shorter of the terminal setae, and three minute setae arranged in a line
up the segment. On the distal half of the outer surface of the outer segment is a
patch of fine spinules. The toothed edge of the coxale has nine unusually elongate
teeth (fig. loD). The distal list has two large teeth, the second of which is very
serrated, followed by 17 small teeth. The proximal list has a large tooth followed
by about six small teeth, another large tooth and a further 20 smaller irregular
teeth. This is in substantial agreement with Rudjakov's (1962) type description.
There is a seta on the distal outer edge of the first segment which Rudjakov noted
as absent but was probably broken off in his specimens.

Maxilla (fig. nC). The maxilla has a long basal segment. There are six anterior,
one lateral and three posterior setae on the first segment. There are no spines

CONCHOECIA FROM THE NORTH ATLANTIC

277

on the outer edge of the segment. The second endopodite segment is long and thin
with long terminal hook setae.

FIG. 10. Conchoecia vitjazi Rudjakov, female. A, carapace right valve mounted flat;
B, frontal organ and first antenna; C, detail of the tip of the frontal organ; D, mandible
tooth lists and toothed edge of the coxale; E, second antenna; F, second antenna endopo-
dite.

2 7 8

M. V. ANGEL

Labrum. The hyaline membrane has a deep rounded notch.

Caudal f urea. The first pair of hook spines just fail to reach level with the ends
of the second pair. There is an unpaired seta dorsal of the hook spines. Between
the lamellae is a covering of fine hairs.

FIG. n. Conchoecia vitjazi Rudjakov, female. A, mandible toothed edge of the basale;
B, mandible endopodite; C, maxilla endopodite.

DISCUSSION

At 'Discovery' station 6665 C. procera Miiller was most abundant by day from
50-300 m, C. microprocera sp. nov. was most abundant at 25 m and occurred in
quite large numbers down to 100 m, C. macroprocera sp. nov. was most abundant

CONCHOECIA FROM THE NORTH ATLANTIC

FIG. 12. Conchoecia decipiens Miiller, male. A, lateral view of the carapace; B, ventral
view of the carapace; C, frontal organ and first antenna; D, frontal organ capitulum;
E, first antenna armature of the b, e and d setae; F, second antenna; G, left second
antenna endopodite; H, right hook appendage; I, left hook appendage; J, sixth limb;
K, penis; L, details of the tip of the penis.

280

M. V. ANGEL

from 500-300 m and was absent from above 200 m. At night C. procera and C.
microprocera showed reverse migrations out of the surface 50-75111, whereas C.
macroprocera did not migrate.

FIG. 13. Conchoecia decipiens Miiller. A, mandible tooth list and toothed edge of the
coxale; B, mandible toothed edge of the basale; C, mandible endopodite; D, maxilla
endopodite; E, labrum; F, fifth limb.

CONCHOECIA FROM THE NORTH ATLANTIC

281

The other species which have been attributed to the 'procera' group are C. decipiens
Miiller igoGa, C. brachyaskos Miiller I9o6a and C. vitjazi Rudjakov 1962. C.
decipiens has only been reported from the Indian Ocean from between 23 N (Leveau
1968) and 25 S (Miiller 1908). To show clearly the differences between C. decipiens
Miiller and the other species, especially C. macroprocera sp. nov. drawings of speci-
mens lent me by Mr J. George of the Cochin Indian Ocean Sorting Centre are in-
cluded (figs 12, 13, 14). Table I also includes measurements of the specimens.

FIG. 14. Conchoecia decipiens Miiller, female. A, lateral view of the carapace; B, ventral
view of the carapace; C, frontal organ and first antenna; D, second antenna; E, second
antenna endopodite; F, male, detail of the posterior dorsal corner of the carapace.

282 M. V. ANGEL

C. decipiens Miiller has a large spine and secondary spine on the right carapace
valve at the posterior dorsal corner. In the males the following characters distin-
guish C. decipiens from C. macroprocera: (i) The capitulum of the frontal organ is
distinctive. (2) The first antenna is relatively longer in C. decipiens. (3) The
first antenna e seta armature has no distally pointing spinules and the main spines
decrease in size proximally. (4) On the second antenna the protopodite is longer
and the shape of the right hook appendage is distinctive. (5) The endopodite of
the mandible carries two setae on the inner face of the first segment. (6) The
penis has five oblique muscles.

In the females the following characters distinguish C. decipiens from C. macro-
procera: (i) The outline of the carapace, (2) the shape of the frontal organ, (3) the
endopodite of the second antenna has no spinules on the first segment near the bases
of the a and b setae, the second segment is completely bare and the g seta is markedly
flattened.

Miiller (igo6a) included C. brachyaskos in the 'procera' group on the basis of the
female frontal organ and the shortness of the sensory setae on the endopodite of
the male second antenna. However, because of the setation of the male first
antenna and its distinctive e seta armature (Miiller igo6a T. XIV fig. 12), it has not
been included here. C. brachyaskos Miiller was present in the deep midwater samples
from 'Discovery' station 6665. There are two size groups; the larger occurring
only from below 1000 m. This large form is superficially very similar to the Ant-
arctic form and so this species requires careful taxonomic investigation before its
status can be certain.

The discovery of a specimen attributable to C. vitjazi Rudjakov in the North
Atlantic provides an interesting example of the faunistic relationships between the
Pacific and the North Atlantic (c.f. Briggs 1970). This species, only recorded
before from below 6000 m in the Kurile-Kamchatka Trench (463i'N, i5422'E,
4348'N, i4955'E), was considered by Rudjakov (1962) to be endemic to that region.
The 'Discovery' specimen does show some minor variations from the type description,
but these are not considered great enough to attribute to it new specific rank. The
male of the species is yet to be described and so the inclusion of the species in the
'procera' group is yet to be confirmed.

SUMMARY

1. Conchoecia procera Miiller is shown to have been confused with two very
closely related but distinct species in the past.

2. C. procera sensu strictu is described and the two new species are described
and named C. microprocera sp. nov. and C. macrorpocera sp. nov.

3. An adult female ascribed to C. vitjazi Rudjakov 1962 is described from the
North Atlantic ; a species previously thought to be endemic to the Kurile-Kamchatka
Trench.

4. The inclusion of C. brachyaskos Miiller in the 'procera' group is questioned.

CONCHOECIA FROM THE NORTH ATLANTIC 283

REFERENCES

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- i96ga. Planktonic ostracods from the Canary Island region; their depth distributions,
diurnal migrations and community structure. /. mar. biol. Ass. U.K. 49 : 515-553.

- ig69b. Repeated samples from a deep midwater planktonic ostracod community. /.
exp. mar. biol. Ecol. 3 : 76-89.

BRIGGS, J. C. 1970. A faunal history of the North Atlantic Ocean. Syst. Zoo/. 19 : 19-34.
CLAUS, C. 1890. Die Gattungen und Arten der mediterranen und atlantischen Halocypriden
nebst Bermerkungen liber die Organisation derselbe. Arb. Zoo/. Inst. Wien 9 : 1-34.

- 1894. Die Halocypriden und ihre Entwicklungsstadien. Gesammelt 1890, 1891, 1892,
1893. Denkschr. Akad. Wiss., Wien Math.-nat. Kl. 61 : i-io.

CURRIE, R. I. & FOXTON, P. 1957- A new quantitative plankton net. J. mar. biol. Ass.

U.K. 36 : 17-32.
DEEVEY, GEORGIANA B. 1968. Pelagic ostracods of the Sargasso Sea off Bermuda. Bull.

Peabody Mus. nat. Hist. 26 : 1-125.
FOWLER, G. H. 1909. The Ostracoda. Biscayan Plankton, XII. Trans. Linn. Soc. Land.,

(2) (Zoo/.) 10 : 219-336.
FOXTON, P. 1963. An automatic opening-closing device for large plankton nets and midwater

trawls. /. mar. biol. Ass. U.K. 43 : 295-308.

- 1969. SOND cruise 1965. Biological sampling methods and procedures. /. mar.
biol. Ass. U.K. 49 : 603-620.

GRANATA, L. & Di CAPORACCIO, L. 1949. Ostracodes marins recueillis pendant les croiseres
du Prince Albert i er der Monaco. Res. Camp. sci. Monaco 109 : 1-51.

HURE, J. 1955. Distribution annuelle verticale du zooplancton sur une station de 1'Adriatique
meridionale. A eta Adriat. 7 : 3-69.

- 1961. Distribution saisonniere et migration journaliere verticale du zooplancton dans la
region profonde de 1'Adriatique. Acta Adriat. 9 : 3-59.

ILES, E. J. 1953. A preliminary report on the Ostracoda of the Benguela Current. 'Dis-
covery' Rep. 57 : 259-280.

LEVEAU, M. 1965. Contribution a 1'etude des Ostracodes et Cladoceres du Golfe de Marseille.
Rec. Trav. St. mar. Endoume 37 : 161-243.

- 1967. Ostracodes pelagiques du sud-ouest de 1'Ocean indien (Regeon de Tulear). Rec.
Trav. St. mar. Endoume, suppl. 6 : 63-70.

- 1969. Ostracodes pelagiques recuellis lors de la 3 e campagne de 1'aviso 'Commandant
Robert Giraud'. Rec. Trav. St. mar. Endoume, suppl. 8 : 127-280.

MULLER, G. W. 1890. Ueber Halocypriden. Zoo/. Jb. Abt. System. 5 : 253-280.

- 1894. Die Ostracoden des Golfes von Neapel und der Angrenzenden Meeres-abschnitte.
Fauna Flora Golf. Neapel 21 : 29-154.

I9o6a. Ostracoda. Wiss. Ergebn. Deutsch. Tiefsee-Exped. 8 : 29-154.

I9o6b. Ostracoden der Siboga Expedition. Siboga Exped. 30 : 1-40.

- 1908. Die Ostracoden der Deutschen Siidpolar Expedition 1901-1903. Sudpolar
Exped. 10 : 51-182.

1912. Ostracoden. Das Tierreich, 31 : 1-434.

RUDJAKOV, J. A. 1962. Ostracoda Myodocopa of the family Halocypridae from the north-
western Pacific. Trudy Inst. Okeanol. 58 : 172-201. (Translation available from N.I.O.)

SCHWEIGER, L. 1912. Adriatische Cladoceren und Plankton-Ostracoden. Sber. Akad. Wiss.
Wein, Math.-Nat. Kl. 121 : 239-272.

Dr M. V. ANGEL

NATIONAL INSTITUTE OF OCEANOGRAPHY

WORMLEY, GODALMING, SURREY

A LIST OF SUPPLEMENTS
TO THE ZOOLOGICAL SERIES

OF THE BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

1. KAY, E. ALISON. Marine Molluscs in the Cuming Collection British Museum
(Natural History) described by William Harper Pease. Pp. 96; 14 Plates.
1965. (Out of Print.) 3.75.

2. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier and
Valenciennes. Pp. 180; n Plates, 15 Text-figures. 1967. 4.

3. TAYLOR, J. D., KENNEDY, W. J. & HALL, A. The Shell Structure of Mineralogy
at the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125; 29 Plates
77 Text-figures. 1969. 4.50.

4. HAYNES, J. R. Cardigan Bay recent Foraminifera (Cruises of the R.V. Antur)
1962-1964. (In press.)

Printed in England by Staples Printers Limited at their Kettering, Northants, establishment

J MISCELLANIA

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 21 No. 8

LONDON : 1972

MISCELLANIA

pp. 285-361

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 21 No. 8

LONDON: 1972

THE BULLETIN OF THE BRITISH MUSEUM

(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical series.

Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
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within one calendar year.

In 1965 a separate supplementary series of longer
papers was instituted, numbered serially for each
Department.

This paper is Vol. 21, No. 8 of the Zoological
series. The abbreviated titles of periodicals cited follow
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Trustees of the British Museum (Natural History), 1972

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Issued 4 February, 1972 Price 2.60

CONTENTS

Page

Conchoecia pseudoparthenoda. By MARTIN VIVIAN ANGEL . . . 289

Genus Lithobius. By EDWARD HENRY EASON 297

The Chestnut-shouldered Wren. By COLIN JAMES OLIVER HARRISON . 313

Lironeca (Isopoda; Cymothoidae). By ROGER JOHN LINCOLN . . . 329

Eunice manihine (Polychaeta; Eunicidae). By MICHAEL ROY LONGBOTTOM 339

The Gangetic Dolphin Platanista gangetica. By GEORGE PILLERI . . 345

A new Genus and Species of Sudan Leech. By LAURENCE REGINALD

RICHARDSON. .......... 349

Genera Phoxinellus, Pseudophoxinus and Paraphoxinus. By ETHELWYNN

TREWAVAS 359

CONCHOECIA PSEUDOPARTHENODA (NOV. SP)

A NEW HALOCYPRID OSTRACOD FOR THE

TROPICAL NORTH ATLANTIC

By M. V. ANGEL

INTRODUCTION

IN the Spring of 1968 a series of horizontal tows were made by R.R.S. Discovery at
station 6665 to study the vertical distribution of planktonic animals at 10 16' N,
19 47' W. A new Conchoecia species belonging to the magna group and very
closely related to C. parthenoda Miiller 1906 occurred in the near surface hauls.
The species is described here and named Conchoecia pseudoparthenoda.

Conchoecia pseudoparthenoda n. sp.

The type specimens were from haul 36 at station 6665 from a depth of 25 m at
1637-1715 hrs on 26 February 1968, position 10 16' N, I947'W. They are
deposited mounted on slides in the British Museum (Natural History), male No.
1971.2.1.5, female No. 1971.2.1.6.

MALE. The range in carapace length of 120 specimens was 1-56-1-72 mm with
a mean of 1-637 '37 mm. The outline of the carapace (Fig. lA) is similar to
that of C. parthenoda Miiller (Deevey 1968, Angel I969a). The position of the left
asymmetrical gland is quite distinctive; in the type specimen it opens 0-48 mm
anterior of the posterior carapace hinge. The right asymmetrical gland opens at
the usual position on the posterior ventral corner. Fine spines occur along the
dorsal surface of the carapace in the region of the hinge line, and on the edges of the
valves from below the rostral incisure to about the midpoint of the ventral side.

Frontal organ. The shaft extends to level with the end of the limb of the first
antenna (Fig. iB). The capitulum is down-turned, its posterior edge is almost
straight and carries spines down the proximal two thirds (Fig. iC). The anterior
edge is slightly concave so that the capitulum is narrowest in its middle region, and
carries a few spines near its base. The end of the capitulum is rounded.

First Antenna. The two basal segments are sub-equal and bare (Fig. iB). The
a seta lies back parallel with the limb reaching level with or just short of the joint
between the two basal segments. The c seta is short. The b seta carries nine fine
spines level with the distal end of the armature on the e seta (Fig. iD), whereas the
d seta carries about 27 fine spines. The b seta is only slightly shorter than the d
seta, which is only just shorter than the e seta. The e seta armature consists of 9
to 10 paired spines followed by 22-26 alternating spines, with a total range of 40-44
spines (Fig. iD). The e seta also has a few scattered spines on its anterior leading
edge close to its base.

Second Antenna. The protopodite is more than half the length of the carapace

Bull. Br. Mus. nat. Hist. (Zool.) 21, 8

2 go

M. V. ANGEL

and three times the first exopodite segment (Fig. lE). All the exopodite segments
are bare. The longest swimming seta is two fifths the carapace length. On the
endopodite, the a seta is bare and curves back behind the b seta (Fig. iG). The b

FIG. i. Conchoecia pseudoparthenoda male. A. Outline of carapace. B. Frontal organ
and first antenna, c. Capitulum of frontal organ. D. Detail of armature of the anten-
nular e, b and d setae. E. Second antenna. F. Copulatory organ. G. Endopodite of
the left second antenna. H. Endopodite of the right second antenna.

CONCHOECIA PSEUDOPARTHENODA (NOV. SP.) 291

seta carries three long hairs near its base and fine spines distally. The processus
mamillaris is bluntly pointed. The c and d setae are sub-equal and rather short.
The e seta is minute. The g seta is almost as long as the protopodite and the f seta
is similar in length to the longest swimming seta. The h, i, and j setae are half the
length of the f seta and about two-fifths the length of the g seta. They have weakly
developed shafts. The right hook appendage is right angled near its base, curves
through a further 90 half way along its length ending in a point with subterminal
ridging (Fig. iH). The left hook appendage is bent through less than a right angle
near its base and is then straight (Fig. iG). Towards its end it narrows and ter-
minates in a curved point with subterminal ridging.

Mandible. The basal segment is long (Fig. zC). The first exopodite segment
carries two setae on its inner edge, one twice the length of the other. The toothed
edge of the pars incisa has two spine teeth followed by six broad finely serrated
teeth, and a single inner broad tooth (Fig. 26). Hairs are inserted near the bases of
the spine teeth and the outer setae. These setae project only just beyond the level
of the teeth. The coxale toothed edge has ten bluntly pointed teeth (Fig. 2 A).
The distal tooth list has two large teeth ; the second is serrated and followed by 13
small blunt teeth. The proximal list has about 14 irregularly sized teeth.

Maxilla. The basal segment carries a seta (Fig. 2D). The first endopodite
segment has six anterior, one lateral and three posterior setae. There is a group of
about six short spines at the end of the segment.

Labrum. This is the usual shape for the magna group (Fig. 2E).

Caudal Furca. The furca has eight pairs of hook spines (Fig. 2F). The first pair
does not reach the level of the ends of the second pair. There is no unpaired post-
erior seta.

Copulatory organ. This has eight oblique muscles (Fig. iF). The terminal edge
which curves in and over the intromittent spine is serrated.

FEMALE. The range in length of 222 specimens was i -72-1 -90 mm with a mean
of 1-841 0-038 mm. As in the male the left asymmetrical gland opens 0-48 mm
anterior of the posterior hinge, and the right gland on the posterior ventral corner
(Fig. 3 A). Fine spines occur on the dorsal surface in the vicinity of the hinge line
and round the ventral edges of the valves in the anterior half. In many specimens
there were concentric striations clearly visible over the whole of the carapace.

Frontal organ. It is straight with no separation into shaft and capitulum (Fig.
36). It terminates in a long downturned point. There are a few spines on its
ventral edge (Fig. 36).

First Antenna. The segmentation of the limb is indistinct. The long spinous
dorsal seta reaches almost level with the tip of the frontal organ (Fig. 36, C). There
is a group of spinules close to the insertion of the e seta. The e seta is twice the
length of the limb and carries spines on its trailing posterior edge distal of the ends
of the other setae. The other four setae are a third the length of the e seta.

Second Antenna. The protopodite is just less than half the carapace length and
nearly three times the length of the first exopodite segment (Fig. 3D). All the
exopodite segments are bare. The longest swimming seta is three quarters the
length of the protopodite. On the endopodite the a seta is half the length of the

292

M. V. ANGEL

b seta, and both are finely spinous (Fig. 3E). The processus mamillaris has a small
pointed tubercle at its tip. The g seta is just over half the length of the protopodite.
It is unflattened and carries fine spinules near its tip. The f seta is only a little
longer than the h, i and j setae which are two thirds the length of the g seta.

B 005

FIG. 2. Conchoecia pseudoparthenoda. A. Toothed edge of coxale and tooth lists of the
mandible. B. Toothed edge of the mandibular basale. c. Endopodite of the mandible.
D. Maxilla endopodite. E. Labrum. F. Caudal furca.

CONCHOECIA PSEUDOPARTHENODA (NOV. SP.)

293

JUVENILES. Two juvenile stages were recognised from the hauls by the position
of the left asymmetrical glands. The smallest group consisting of 106 specimens
ranged in size from 0-86-0-96 mm with a mean of 0-903 + 0-022 mm. The final
stage juveniles ranged in size from 1-22-1-34 mm with a mean of 1-286 + 0-026 mm
for 163 specimens.

COMPARISON BETWEEN C. pseudoparthenoda AND C. parthenoda. In the females,
C. pseudoparthenoda often has more distinctive sculpturing and the posterior ventral
region of the carapace is more strongly developed. In both sexes the left asym-

FIG. 3. Conchoecia pseudoparthenoda female. A. Outline of carapace. B. Frontal organ
and first antenna, c. Details of frontal organ and the end of the first antenna. D. Second
antenna. E. Endopodite of the second antenna.

294 M - v - ANGEL

metrical gland is 0-48 mm anterior of the posterior hinge, and this is diagnostic of
the species; in C. parthenoda the gland opens 0-24 mm anterior of the posterior
hinge. The carapace breadth is less in C. pseudoparthenoda. The shapes of the
frontal organs of the females of the two species are quite distinct. The dorsal seta
of the first antenna is shorter in the female of C. parthenoda and this species has no
spinules at the insertion of the first antennal e seta, also the e seta is relatively
longer. Similarly the exopodite segments and the endopodite setae of the second
antennae of the females are all relatively longer in C. parthenoda. A comparison
of the lengths of the various setae and other meristic characters of the two species
are shown in Table i.

In the males C. parthenoda has a longer b seta on the first antenna and a shorter
c seta ; the b and d setae carry more spines. The antennular e seta armature usually
consists in both species of nine pairs of distal spines. In C. parthenoda the paired
spines are followed by about 18 alternating spines; giving a total range of 35-38
spines. In C. pseudoparthenoda the paired spines are followed by about 24 alter-
nating spines ; giving a total range of 40-44 spines.

On the mandibles the proximal tooth lists are quite distinct ; C . parthenoda having
three large teeth and 16 smaller teeth, and C. pseudoparthenoda 14 irregular teeth.
On the caudal furca C. parthenoda has an unpaired dorsal seta but C. pseudopar-
thenoda has none. The live colouration of the two species is similar, for the most
part they are transparent and colourless or very pale red. The anterior ventral
edges of the carapace valves are yellow, and the pharyngeal region is yellow or
orange.

Bioluminescence has not been observed in either of the two species.

DISCUSSION

At station 6665 C. pseudoparthenoda predominated over C. parthenoda at 25 m,
but C. parthenoda was the more numerous in the 50 m and 75 m hauls. Both
species were virtually restricted to the surface 100 m. C. pseudoparthenoda was
absent from the plankton off Fuerteventura (Canary Islands) and the Moroccan
coast (Angel 1968, 1969^, but Deevey (personal communication) has found what is
probably this species to be a rare member of the plankton off Bermuda. C. par-
thenoda was common in all these areas.

Poulsen (1969) has recently described a male specimen from the Congo region
with the gland openings of the left asymmetrical gland 0-38 mm from the posterior
dorsal hinge (see his figure 10 and table p. 154). The females, however, had gland
openings 0-27 mm from the hinge. The carapace lengths of all his specimens were
slightly larger than described here for C. pseudoparthenoda, and appreciably longer
than for C. parthenoda. His description of the male antennular e seta armature
makes no mention of the distal spines being paired, and the total counts are con-
siderably lower than for either C. parthenoda or C. pseudoparthenoda. It seems
unlikely from his description that Poulsen's specimens belonged to either of these
species.

C. pseudoparthenoda appears to have an equatorial distribution in the Atlantic
and may prove to be endemic to this region.

CONCHOECIA PSEUDOPARTHENODA (NOV. SP.)

295

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REFERENCES

ANGEL, M. V. 1968. The thermocline as an ecological boundary. Sarsia 34 : 299-312.

- igGga. The redescription of three halocyprid ostracods, Conchoecia hyalophyllum Claus,
C. magna Claus and C. parthenoda Mullet from the North Atlantic. Crustaceana 17, pp.

45-63-

- 19695. Planktonic ostracods from the Canary Island region: their depth distributions,
diurnal migrations and community organization. /. mar. biol. Ass. U.K. 49 : 515-553.

DEEVEY, Georgiana. 1968. Pelagic ostracods of the Sargasso Sea off Bermuda. Bull.

Peabody Mus. nat. Hist. 26 : 1-125.

MULLER, G. W. 1906. Ostracoda. Wiss. Ergebn. Deutsch. Tiefsee-Exped. 8 (2), pp. 29-154.
POULSEN, E. M. 1969. Ostracoda-Myodocopa from the Eastern Tropical Atlantic. Vidensk.

Meddr. dansk naturh. Foren. 132 : 129-197.

Dr. M. V. ANGEL

NATIONAL INSTITUTE OF OCEANOGRAPHY

WORMLEY,

GODALMING, SURREY

THE TYPE SPECIMENS AND IDENTITY OF THE

SPECIES DESCRIBED IN THE GENUS

LITHOBIUS BY GEORGE NEWPORT IN 1844,

1845 AND 1849 (CHILOPODA, LITHOBIOMORPHA)

By E. H. EASON

INTRODUCTION

GEORGE NEWPORT described eighteen nominal species under Lithobius one of which,
L. emarginatus, he later removed to the genus Henicops. Pocock (1890, iSgia,
i8gib, 1901) examined the type specimens of at least eight of these species but he
only identified five, one of them incorrectly. Of the remainder, some have been
identified more or less definitely by various authors from their original descriptions
but the identity of the others has only been tentatively suggested or has, hitherto,
been quite unknown. The type specimens of sixteen of Newport's species are
preserved either in the British Museum (Natural History), the Hope Department of
Zoology, Oxford or the Museum National d'Histoire Naturelle, Paris. They have
all been re-examined for the purpose of the present study and their identity is either
confirmed or established for the first time. It is not always possible to tell whether
Newport had one or several specimens before him when he wrote his descriptions and
he made no formal designation of type specimens, but where only a single specimen
is available it is regarded as the holotype: otherwise a lectotype is selected where
necessary. An attempt is also made to determine the identity of the species for
which type specimens have not been found.

Newport's 1844-45 paper was divided between parts 3 and 4 of volume 19 of the
Transactions of the Linnean Society of London. The section in part 3, which includes
a figure with the caption 'Lateral view of the head of Lithobius americanus, Newp.'
(Tab. 33), was published in November 1844, but the section in part 4, which includes
the written descriptions of this and eight other new species of Lithobius, was not
published until November 1845 (Raphael, 1970). The species described in this paper
are usually dated 1844, but clearly all except L. americanus should be dated 1845.

Conclusions as to the status and present classification of all the nominal species
described by Newport in the genus Lithobius are summarized in Table i.

i. Lithobius hardwickei Newport
Fig. i

Lithobius Hardwickei Newport, 1844, p. 96; 1845 (1844-45), p. 366

TYPE LOCALITY. Singapore.

TYPE SPECIMEN. Holotype : a male pseudomaturus of L.forficatus (Linn.) 16 -5 mm
long, dried and pinned, labelled "L. Hardwickei Newp." in Newport's hand and

Bull. Br. Mus. nat. Hist. (Zool.) 21, 8

298 E. H. EASON

"Hardwicke Bequest, Singapore" on a separate ticket. British Museum (Natural
History).

REMARKS. As Newport stated, this specimen is smaller than an adult of L.
forficatus and of a paler colour than usual : it has only 39 antennal articles (Newport
gave 41) but the prosternum is exactly as described by Newport with 5+8 teeth

(Fig. i).

According to Haase (1887), Pocock examined the specimen and found it to have
posterior projections on T. 9, n and 13 but came to no conclusion as to its identity.
Verhoeff (1937) merely noted that Newport's description of L. hardwickei agreed
with none of the species of Lithobius he found in the Malay peninsula. Wang and
Tang (1965), in the most recently published list of Chilopoda from Singapore, made
no mention of the species and there is little doubt that L. forficatus was introduced
to Singapore but never became established.

2. Lithobius leachi Newport

Lithobius forficatus : Leach, 1814, p. 408

Lithobius Leachii Newport, 1844, p. 96; 1844 (1844-45), Tab. 33, fig. 30

Lithobius sp. Newport, 1844 (1844-45), Tab. 33, fig. 31

Lithobius Leachii: Newport, 1845 (1844-45), p. 368

TYPE LOCALITY. Europe.

TYPE SPECIMEN. Holotype: a female of L. forficatus (Linn.) 20 mm long, dried
and gummed to a card over the whole of its ventral aspect, labelled "L. Leachii
Newp." in Newport's hand. British Museum (Natural History).

REMARKS. Newport examined the original Linnean specimen of L. forficatus and
finding it to differ slightly in the form of the prosternum from Leach's specimen of
this species, referred the latter to a new species, L. leachi. Synonymy of L. leachi
with L. forficatus was tentatively suggested by Meinert (1868) and has never been
disputed. Although the ventral aspect of the holotype is obscured owing to its
being gummed to a card, it is undoubtedly the specimen which Newport examined
and which Leach had, quite correctly, identified as L. forficatus.

3. Lithobius pilicornis Newport

Lithobius pilicornis Newport, 1844, p. 96

Lithobius sp. Newport, 1844 (1844-45), Tab. 33, fig. 34

Lithobius pilicornis : Newport, 1845 (1844-45), p. 369

TYPE LOCALITY. England.

TYPE SPECIMEN. Holotype: a male 26 mm long, dried and pinned, labelled "L.
pilicornis Newp." in Newport's hand. British Museum (Natural History).

REMARKS. This specimen was described by Pocock (iSgia) who gave a full
account of the species.

Gervais, in his list of published descriptions of species of Myriapoda (Walckenaer
and Gervais, 1847), mentioned Lithobius pulchricornis and L. pilicornis as having

GENUS LITHOBIUS

299

been described by Newport in 1844 and 1845 respectively. It seems that pulchri-
cornis attaches to Newport's first (1844) brief description of L. pilicornis. The
name does not appear in any of Newport's works and must have arisen from some
carelessly written transcript of his earlier paper.

FIGS 1-5. i, Lithobius hardwickei. Dental margin of prosternum of holotype, ventral.
3, Lithobius americanus. Dental margin of prosternum of holotype, ventral. 2, Litho-
bius brevicornis. Dental margin of prosternum of holotype, ventral. 4, Lithobius
elongatus. gth to i4th tergites of lectotype, dorsal. 5, Lithobius monilicornis. gth to
1 4th tergites of male syntype, dorsal.

3 oo E. H. EASON

4. Lithobius sloanei Newport

Lithobius Sloanei Newport, 1844, p. 96; 1845 (1844-45), p. 369
TYPE LOCALITY. Unknown.

TYPE SPECIMEN. Holotype: a bleached and distorted female of L. pilicornis
about 30 mm long, dried and pinned, labelled "L. Sloanei Newp." in Newport's
hand and "Sir H. Sloane's coll. ?4i67" on a separate ticket. British Museum
(Natural History).

REMARKS. This specimen was described and discussed by Pocock (iSgia) who
identified it correctly and noted that the number 4167 corresponds to the following
entry in Sir Hans Sloane's catalogue "a middling good sized brown Scolopendra".

5. Lithobius castaneus Newport

Lithobius castaneus Newport, 1844, p. 96; 1845 (1844-45), p. 370

TYPE LOCALITY. Sicily.

TYPE SPECIMEN. Holotype: a female 20 mm long, dried and pinned, labelled "L.
castaneus Newp." in Newport's hand. British Museum (Natural History).

REMARKS. This specimen was examined by Pocock (1890) who recognized its
identity with L. eximius Meinert, 1872. He noted correctly that it only has 26
antennal articles (Newport gave 41). It agrees with Brolemann's (1930) description
of L. castaneus except for the presence of the spine 15 VaT which was recorded by
Meinert for L. eximius, and the extension of the distinctive sculpturing of the large
tergites, rather faintly, on to T. 8, 10 and 12 : Brolemann described this last feature
on T. 1.3 and 5 only. Other characters possessed by the holotype, all in agreement
with Brolemann's description, are 6, 7, 7, 7 oblong coxal pores, lateral spines on the
i4th and I5th coxae, and a simple claw on the gonopod with only a feeble lobe at the
base of its external ridge and no denticles. These features place it in the subspecies
L. castaneus buchnerorum described from Ischia by Verhoeff (1942) who assumed that
the typical form of the species has a dentate female genital claw (Verhoeff, 1934).
However, as Matic (1961) pointed out, many of the subspecies of L. castaneus
described by Verhoeff were based on unstable characters and are probably without
validity : should it be found that a geographical race with a dentate genital claw can
be defined it can not be regarded as belonging to the nominate subspecies.

6. Lithobius emarginatus Newport

Lithobius emarginatus Newport, 1844, p. 96

Henicops emarginatus: Newport, 1845 (1844-45), p. 372

TYPE LOCALITY. New Zealand.

TYPE SPECIMEN. Holotype: a very defective female 8-5 mm long, labelled "H.
emarginatus Newp." and "in the ground Capt. Ross" in Newport's hand: the speci-
men has been pinned through T. 7 damaging this and the adjacent tergites, but the
pin has been withdrawn and the specimen is gummed to the apex of a triangular
piece of card. British Museum (Natural History).

GENUS LITHOBIUS

301

REMARKS. Pocock examined this specimen, redescribed the genera Henicops
Newport and Lamyctes Meinert, and removed emarginatus to the latter (Pocock,
1901). Archey (1937) also saw the specimen which he described as being 'much
shrivelled and somewhat mutilated'. He noted however that the 2+2 prosternal
teeth were discernible in the type and it is confirmed that in this as in all other
respects the specimen is identical with the common New Zealand species Archey
described under Lamyctes emarginatus.

7. Lithobius rubriceps Newport

Lithobius rubriceps Newport, 1845 (1844-45), p. 364
TYPE LOCALITY. Southern Spain.

REMARKS. Although Newport deposited his material belonging to this species in
the British Museum, no type specimen has been found. It seems significant that
Pocock, who examined and reported on most of Newport's type specimens of
Lithobius in the British Museum, made no mention of L. rubriceps in any of his
writings: this suggests that the specimen had already been lost or badly damaged
before 1890 when Pocock made his first observations.

However, the space in the cabinet containing Newport's specimens and apparently
allotted to L. rubriceps is occupied by four examples of the common Iberian species
known as L. insignis Meinert, labelled "Lithobius rubriceps Newport, Lisbon" and
"96.3.8.103-106, Pascoe". These specimens were collected by the entomologist
F. P. Pascoe and presented to the Museum by Miss Pascoe. They are accompanied
by two further specimens of L. insignis labelled "Cintra 25.11.96". Newport's
description is unmistakable and there is no doubt that these specimens were correctly
identified, probably by Pocock, and that L. rubriceps is the senior synonym of
L. insignis.

Meinert (1872) described L. insignis and L. gracilipes as new species from Spain,
suggesting L. rubriceps as a possible synonym of the latter. This choice on the part
of Meinert is difficult to understand because Newport's description of rubriceps is
much closer to insignis than to gracilipes. However, gracilipes only differs from
insignis in being smaller with fewer coxal pores and may well prove to be another
synonym of L. rubriceps.

The species was fully described by Machado (1952).

8. Lithobius fasciatus Newport

Lithobius fasciatus Newport, 1845 (1844-45), p. 365
TYPE LOCALITY. Florence and Naples.

TYPE SPECIMENS. Lectotype: a specimen labelled "Lithobius fasciatus Newp." in
Newport's hand. Paralectotypes: two unlabelled specimens accompanying the
lectotype. Hope Department of Zoology.

REMARKS. These three specimens have been fully described in an earlier paper
(Eason, 1970). They were examined by Pocock (1890) who was mistaken in
believing them to be identical with L. grossipes C. L. Koch.

302 E. H. EASON

9. Lithobius multidentatus Newport

Lithobius multidentatus Newport, 1845 (1844-45), p. 365
TYPE LOCALITY. New York.

TYPE SPECIMEN. Holotype: a female 22 mm long labelled as the holotype of
Lithobius multidentatus Newport by Dr. R. E. Crabill who removed it from the
cabinet containing Newport's dried specimens in 1960, relaxed it in trisodium
phosphate and placed it in spirit. British Museum (Natural History).

REMARKS. Wood (1865) recognized the identity of L. multidentatus with the
species he had himself described under Bothropolys nobilis (Wood, 1863), apparently
basing his conclusion on Newport's very brief description. In fact, the holotype has
9 + 9 prosternal teeth and not 8 + 8 as Newport stated. The specimen does,
however, agree in all respects with Chamberlin's (1925) detailed description of
Bothropolys rmdtidentatus except for the presence of a small extra medial spur on
the left gonopod, in addition to the usual two.

It is clear from Dr. Crabill's labelling of the holotype that he was satisfied as to its
identity with the common North American species generally known as B. multi-
dentatus.

10. Lithobius americanus Newport

Fig. 3
Lithobius americanus Newport, 1844 (1844-45), Tab. 33, fig. 29; 1845 (1844-45), p. 365

TYPE LOCALITY. North America.

TYPE SPECIMEN. Holotype: a rather defective male of L. forficatus (Linn.) 24 mm
long, dried and pinned, labelled "Lithobius Americanus Newp. N.S." in Newport's
hand. Hope Department of Zoology.

REMARKS. Newport distinguished this specimen from L. forficatus by its larger
size and the form of the prosternum. The teeth of the latter are, indeed, irregularly
spaced (Fig. 3) as in so many examples of L. forficatus.

Synonymy of L. americanus with L. forficatus was first proposed by Stuxberg
(1871) and has never been disputed.

ii. Lithobius planus Newport

Lithobius sp. Newport, 1844 (1844-45), Tab. 33, fig. 32
Lithobius planus Newport, 1845 (1844-45), p. 366

TYPE LOCALITY. North America.

TYPE SPECIMEN. Holotype: a rather defective male of Bothropolys multidentatus
18 mm long, dried and pinned, labelled "Lithobius planus Newp." in Newport's
hand. Hope Department of Zoology.

REMARKS. Newport made no use of the arrangement of the coxal pores in his
system and so failed to detect the affinity between L. planus, which he believed to be
close to L. variegatus Leach, and L. multidentatus. Further, the holotype of planus

GENUS LITHOBIUS 303

has 7 -f 7 prosternal teeth whereas multidentatus was described as having 8 +8 which
is the more usual number in this species. Wood (1863, 1865) mentioned L. planus
but merely reiterated Newport's description and made no suggestion as to its
identity with any North American species known to him.

12. Lithobius argus Newport

Lithobius Argus Newport, 1845 (1844-45), p. 369
TYPE LOCALITY. Wellington, New Zealand.

TYPE SPECIMENS. Syntypes: a female pseudomaturus 18 mm long and a male
praematurus 12-5 mm long, both of L. forficatus (Linn.), dried and pinned, labelled
"Lithobius Zelandicus Newp." in Newport's hand. Hope Department of Zoology.

REMARKS. Newport distinguished this form from L. forficatus by its smaller
size ; only the female answers closely to his description.

Pocock (iSgib) examined these specimens, noted that they were labelled "Litho-
bius Zelandicus" , and confirmed that they belonged to Lithobius s.s. but came to no
conclusion as to their exact identity. It seems that Newport first named them
Zelandicus and then changed the name to Argus when writing his paper.

This species has never been rediscovered in New Zealand but Archey (1937), in
his most recently published account of the Chilopoda of that country, never ques-
tioned its validity and regarded Newport's record as evidence of the occurrence of
an indigenous species of Lithobius in a country where the Chilopod fauna consists
otherwise almost entirely of genera confined to the southern hemisphere. There is
no doubt that L. forficatus was introduced to New Zealand but lack of any subsequent
records suggests that it has never become established.

13. Lithobius brevicornis Newport
Fig. 2

Lithobius brevicornis Newport, 1845 (1844-45), p. 370

TYPE LOCALITY. Naples.

TYPE SPECIMEN. Holotype : a male pseudomaturus of L. forficatus (Linn.) 17 mm
long, dried and pinned, labelled "Lithobius brevicornis Newp." in Newport's hand.
Hope Department of Zoology.

REMARKS. This specimen has 43 antennal articles, 15 ocelli on each side and
4 +6 prosternal teeth (Fig. 2) , whereas Newport gave 41 articles, 20 ocelli and 6 -f-6
teeth. There is, however, no suggestion in the description that it was based on more
than one specimen and this description seems, therefore, to have been inaccurate.
Newport equated L. brevicornis with L. vesuvianus Costa, but no account of the latter
appears ever to have been published.

Fanzago's (1874) and Fedrizzi's (1877) accounts of L. brevicornis are mere reitera-
tion of the original description and Fedrizzi's mention of the antennae as having 14
articles was obviously due to a misprint. Although L. brevicornis figures in a

304 E. H. EASON

number of more recent Italian faunal lists it has never been redescribed and no
suggestion has ever been made as to its identity.

14. Lithobius melanops Newport

Lithobius melanops Newport, 1845 (1844-45), p. 371
TYPE LOCALITY. Sandwich, England.

TYPE SPECIMEN. Holotype : a very defective female 13 mm long, dried and pinned,
labelled "Lithobius melanops Newp. (Kent 44.41)" in Newport's hand. British
Museum (Natural History).

REMARKS. Pocock (1890) examined this specimen and concluded that it was
conspecific with a common European species widely referred to as L. glabratus C. L.
Koch 1847. Although the antennae and most of the legs are missing there is no
doubt that Pocock was correct.

The species was described fully by Brolemann (1930).

15. Lithobius platypus Newport

Lithobius platypus Newport, 1845 (1844-45), p. 371

TYPE LOCALITY. Egypt.

REMARKS. On the single plate devoted to Myriapoda and Hexapoda Aptera in
the Natural History portion of Savigny's "Description de 1'Egypte", figure 3
represents a small lithobiomorph centipede, 11-5 mm long with short antennae each
of 20 articles, i +2, i ocelli on each side, no posterior tergal projections and with
both i4th and I5th legs thickened. Newport based his description of L. platypus
on this figure which, with the thickened posterior legs, was unlike any of the larger
species of Lithobius with which he was familiar. Although, like all Savigny's
drawings, this figure is beautifully executed it is inaccurate in that a short tergite is
shown immediately behind T. 7; such a tergite is never found in Lithobius or any
related genus. On the other hand, apart from this spurious tergite, the figure is a
very fair representation of L. vosseleri Verhoeff, originally described by Verhoeff
(1901) from Cyprus and recorded by Silvestri (1929) from a number of localities in
North Africa including Alexandria (Egypt). Silvestri referred his specimens to a
new variety, propitia, on the basis of some quite trivial characters and suggested the
possibility of their being identical with L. platypus, the only other species of Lithobius
ever recorded from Egypt.

Savigny's sight failed after he had completed his drawings but before he had
finished his manuscript (Sherborn, 1897), so that no names, descriptions or exact
localities attaching to his specimens of Myriapoda were ever published. Gervais
(1837) commented on Savigny's figure of a lithobiid, but he did not give it a Latin
binomen. Newport's description of L. platypus may, therefore, be the earliest
published description of L. vosseleri, but there is no certainty of this and L. platypus
should be rejected as a nomen dubium.

GENUS LITHOBIUS 305

16. Lithobius elongatus Newport
Fig. 4

Lithobius elongatus Newport in Lucas, 1849, p. 383, PI. 3, figs. 2, 2a, 2b, ac and 2d

TYPE LOCALITY. Lac Tonga, Lac Houbeira and La Calle, Algeria.

TYPE SPECIMENS. Lectotype: a male 24 mm long, preserved in spirit, labelled
"Lithobius elongatus Newport, Algerie Lucas" is here formally designated as the
lectotype. Paralectotypes : a distorted male about 28 mm long and two males 19 mm
long accompanying the lectotype but now placed in a separate tube. Museum
National d'Histoire Naturelle.

REMARKS. Newport's description of L. elongatus is inadequate by modern
standards but this species was more fully described by Verhoeff (1891) under
Lithobius (Polybothrus) koenigi and by Silvestri (1896), in both instances from
Tunisian specimens agreeing in all essentials with the lectotype. Later Silvestri
(1897) described more specimens from Sicily and proposed elongatus and koenigi,
together with L. impressus C. L. Koch and L. monilicornis Newport, as junior
synonyms of L. nudicornis Gervais. L. nudicornis, however, is based on an
extremely scanty description of a specimen from Sicily (Gervais, 1837) an d the name
has been rejected by most authors in favour of elongatus. Whether impressus and
monilicornis, which are usually accepted as synonyms of elongatus, should in fact be
regarded as such is open to question and will be discussed under the next species.
Owing to the uncertainty as to the relative status, not only of elongatus and
impressus (=monilicornis) but of the numerous subspecies which have been des-
cribed from time to time, the lectotype of L. elongatus is described below.

DESCRIPTION OF LECTOTYPE. Size: 24 mm long, 2.75 mm broad at T. 10, i5th
legs 8 mm long. Colour: bleached with little pigmentation. Antennae: 10 mm
long with 44 articles. Ocelli: apparently 1+4, 4, 3 but difficult to see owing to
bleaching. Prosternum : with 5 +5 faintly pigmented teeth ; lateral spines appearing
as unpigmented nodes, each surmounted by a transparent peg, but before bleaching
occurred the lateral spines would, like the teeth, have been strongly pigmented and
would have been counted as teeth by Newport. Tergites (Fig. 4) : posterior angles
of T. 9 right-angled, those of T. n and 13 with projections; T. 10 and 12 longer than
broad with rounded posterior angles. Coxal pores: numerous with no regular
arrangement.

Spinulation :

Ventral Dorsal

CtPFT CtPFT

14 a m amp amp a a amp p p

15 a m amp am a a mp p

i^th prefemur: slightly expanded distally without any conspicuous setal tuft.
i$th prefemur: with an ill-defined dorsal longitudinal sulcus and a moderate internal
distal expansion bearing numerous setae; DpP placed immediately distal to the
setae. i$th apical claw : simple. Gonopods : short and inconspicuous.

306 E. H. EASON

FURTHER SPECIMENS. In addition to the lectotype and three paralectotypes in
the Museum d'Histoire Naturelle there are five males and two females, dried and
pinned, in the British Museum (Natural History) labelled "Lithobius elongatus
Newp." in Newport's hand and "Tunis, 46, 103" on a separate ticket, all of which
answer essentially to the above description. The same may be said of a female
from Tunis (B.M.(N.H.) Reg. no. 90.12.16.23) preserved in spirit along with other
specimens of "L. impressus" collected by Dr. Anderson in North Africa and reported
on by Pocock (1892).

None of these specimens has posterior projections on T. 9 and the number of their
antennal articles varies from 39 to 44. Of variations in spinulation, i4VpT, i4VmT
and J-5"VpF may be present; i5VaF, i5VaT, i5DpP and i5DpF may be absent.
Neither i5DaP nor ventral tarsal spines, both of which are found in the next species,
were present in any of the above specimens.

17. Lithobius monilicornis Newport
Fig- 5

Lithobius monilicornis Newport in Lucas, 1849, p. 384, PL 3, figs 3, 3a, 3b, 30 and 3d

TYPE LOCALITY. Boudjare"a, near Algiers.

TYPE SPECIMENS. Syntypes : a male 30 mm long, a rather defective female 35 mm
long and an immature female 16.5 mm long, all of Eupolybothous impressus (C. L.
Koch), preserved in spirit, labelled "Lithobius monilicornis Newport, Algerie
Lucas". Museum National d'Histoire Naturelle.

REMARKS. Newport distinguished this species from L. elongatus by its more
numerous antennal articles, smaller prosternal teeth, relatively broader T. 8, 10
and 12, and longer legs. All these features are shown by the above specimens which
are undoubtedly identical with the form described from Oran and Algiers by L.
Koch (1862) under L. impressus C. L. Koch, and we have no reason to doubt their
identity with L. impressus as originally described from the Algerian coast by C. L.
Koch (1841) and later figured by the same author (C. L. Koch, 1863).

More distinctive than the characters given by Newport for distinguishing this
form from L. elongatus are the posterior projections on T.g (Fig. 5) and the presence
of ventral tarsal spines on the I4th and i5th legs, both of which were mentioned by
L. Koch in his description of L. impressus.

Synonymy of L. monilicornis with L. impressus was first tentatively suggested by
Meinert (1872).

DESCRIPTION OF MALE. Size: 30 mm long, 4 mm broad at T. 10, i5th legs 12 mm
long; if it were not for the contraction of the trunk (Fig. 5) contrasting with the
extension of that of the lectotype of L. elongatus with exposure of the intersegmental
membranes (Fig. 4), the difference in length between these two specimens would be
more marked. Colour: more bleached than the lectotype of elongatus with prac-
tically no pigmentation. Antennae: 15 mm long with 42 and 43 articles; they
appear to have been damaged and to have undergone imperfect regeneration; the
adult female has 49 and 52 antennal articles. Ocelli: not seen owing to bleaching.

GENUS LITHOBIUS 307

Prosternum: with 6+6 teeth, faintly pigmented at their apices; being more numerous
than in elongatus they appear relatively smaller; lateral spines appearing as unpig-
mented nodes but before bleaching occurred they would, like the teeth, have been
strongly pigmented and would have been counted as teeth by Newport. Tergites
(Fig. 5) : posterior angles of T. 9 with small but distinct projections, those of T.
ii and 13 with more marked projections; T. 10 and 12 broader than long with poster-
ior angles blunt, not rounded. Coxal pores : numerous with no regular arrangement.

Spinulation :

Ventral Dorsal

CtPFTTa CtPFT

14 a m amp amp am a a amp p p

15 a m amp am a a a am p

14^ prefemur: slightly expanded distally without any conspicuous setal tuft.
i$th prefemur: with a well-marked dorsal longitudinal sulcus and a prominent
internal distal expansion bearing numerous setae and a distinct node. i$th apical
claw : simple. Gonopods : short and inconspicuous.

FURTHER SPECIMENS. In addition to the three syntypes in the Museum d'Histoire
Naturelle a female, dried and pinned, in the British Museum (Natural History)
labelled "L. impressus C. Koch" and "96.3.8.107 Pascoe, Oran" and a number of
specimens from various localities in Algeria (B.M.(N.H.) Reg. nos. 90.12.16.20-22
and 24-28; 91.1.18.1-3), preserved in spirit and constituting the majority of the
specimens of "L. impressus" collected by Dr. Anderson in North Africa and
reported on by Pocock (1892), all answer essentially to the above description. They
all have posterior projections on T. 9, the number of their antennal articles varies
from 43 to 50 and in addition to possessing ventral tarsal spines on the I5th and
usually also the I4th legs, 15 DaP is always present whereas 15 DpP is found only
in females.

DISCUSSION. If the specimens mentioned so far in this paper under L. elongatus
(see p. 208) and L. monilicornis ( impressus) were the only ones available for examina-
tion one would have little hesitation in referring the two forms to distinct subspecies
or even to distinct species, but of Dr. Anderson's North African specimens of "L.
impressus", those from Constantine (B.M.(N.H.) Reg. no. 91.1.18.4-7) are inter-
mediate between the two both as regards the shape of the tergites and spinulation,
so it seems that the characters separating elongatus and impressus are unstable.
Neither Polybothrus elongatus nor P. elongatus koenigi as briefly described by
Brolemann (1932) from North Africa can definitely be referred either to elongatus or
to impressus as defined in the present study, but Lithobius (Polybothrus) elongatus
var. oraniensis described by Verhoeff (1901) from a number of localities in the
neighbourhood of Oran seems definitely to belong to L. impressus. L. (Polybothrus)
impressus corsicus Leger and Duboscq (1903), reported from Corsica and the Mari-
time Alps (Brolemann, 1930), agrees with impressus in having posterior projections
on T. 9 and ventral tarsal spines on the i5th legs, but is distinguished by i4DpP
being replaced by a hook. Of the other six described subspecies, all either from the
western Mediterranean islands or the European mainland, none can be easily placed.

308 E. H. EASON

The species or species group to which all these forms belong is clearly in need of
revision but, in spite of the intermediate examples from Constantine, it seems
advisable to retain, for the time being, the distinction between elongatus and
impressus which belong to the genus Eupolybothrus Verhoeff and the subgenus
Allopolybothrus Verhoeff as amended by Jeekel (1967) ; but they should be regarded
as only subspecifically distinct.

The respective ranges of these two subspecies, if indeed they are true subspecies,
are difficult to define exactly. C. L. Koch's figure (C. L. Koch, 1963: fig. 105) is
almost certainly of E. impressus impressus but it was drawn from a specimen from
Bone which is only 80 Km west of La Calle near the Tunisian border, one of the type
localities of E. impressus elongatus. The latter, however, seems to be confined in
North Africa to Tunisia and eastern Algeria whereas E. i. impressus extends along
the Algerian coast westwards to Oran and probably further into Spanish Morocco.

18. Lithobius lucasi Newport

Lithobius Lucasi Newport in Lucas, 1849, p. 385, PI. 3, figs i, la, ib, ic and id

TYPE LOCALITY. Lac Tonga and Lac Houbeira, Algeria.

TYPE SPECIMENS. Syntypes: a male 28 mm long, a female 23 mm long and an
immature male 17.5 mm long, all of L. castaneus, preserved in spirit, labelled
"Lithobius Lucasii Newport, Algerie Lucas". Museum National d'Histoire
Naturelle.

REMARKS. Newport's mention of 5 +5 prosternal teeth, the only character
appearing to distinguish L. lucasi from L. castaneus, is hardly surprising because,
although the stout lateral spines which Newport, no doubt, regarded as teeth are
quite distinct in all three of the above specimens, the 2+2 small teeth are only
evident in the immature male : in both the adult syntypes the dental margin of the
prosternum is twisted and distorted giving the impression of a series of projections
which might be mistaken for teeth. As in the holotype of L. castaneus, the type
specimens of L. lucasi have oblong coxal pores, lateral spines on the I4th and I5th
coxae, and the female has a simple claw on the gonopod with only a feeble lobe at the
base of its external ridge and no denticles: these specimens, therefore, answer to
Verhoeff's (1942) description of L. castaneus buchnerorum.

Brolemann (1921) suggested L. lucasi as a possible synonym of L. castaneus: he
was probably guided by Newport's figure (1849: PI. 3, fig. i) which leaves little
doubt as to its identity.

ACKNOWLEDGEMENTS

My sincere thanks are due to Dr. J. P. Harding and Dr. J. G. Sheals for their
helpful criticism of my manuscript ; to the Trustees of the British Museum (Natural
History) for facilities for studying material; to Mr. K. H. Hyatt of the Arachnida
Section of the Department of Zoology of the Museum for his personal help; to
Professor G. C. Varley and Mr. E Taylor of the Hope Department of Zoology,
Oxford, for enabling me to examine specimens from the Newport Collection ; and to
the Museum National d'Histoire Naturelle, Paris and M. J.-M. Demange for lending
me the type specimens of Newport's Algerian species.

GENUS LITHOBIUS

309

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REFERENCES

ARCHEY, G. 1937. Revision of the Chilopoda of New Zealand. Rec. Auckland Inst. Mus.

2 : 43-100.
BROLEMANN, H. W. 1921. Liste des Myriapodes signales dans le nord de 1'Afrique. Bull.

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E. H. EASON

BOURTON FAR HILL FARM

MORETON-IN-MARSH

GLOS.

A RE-EXAMINATION OF THE CHESTNUT-
SHOULDERED WREN COMPLEX OF AUSTRALIA

By C. J. O. HARRISON

SYNOPSIS

THE chestnut-shouldered wren complex within the genus Malurus is re-examined, using recently
collected material. Seven forms are recognized elegans, pulcherrimus , lamberti, assimilis
(including master si and bernieri), rogersi, dulcis and amabilis. Variations occur within assimilis,
rogersi and amabilis. Some variant individuals from the northern edge of the range of assimilis
show aspects of plumage colour approaching those of other nearby forms. The type of distribu-
tion appears to provide an example of the refuge concept suggested by Keast. It is suggested
that pulcherrimus originated in the Eyre peninsula region and assimilis in the Hamersley region,
and that adaptation to warmer and drier conditions enabled these forms to spread with sub-
sequent climatic amelioration. A wide tolerance of habitat is shown by assimilis. Other
forms appear to have more specific preferences but may be occupying the ecological equivalent,
within their range, of the general habitat required. Evidence of interaction and difference
between forms indicates that elegans and pulcherrimus behave as good species.

The taxonomic status of the other five forms appears to be equal, but whether this should be
specific or subspecific must remain undecided until there is more information on distribution
and possible interbreeding.

Certain broad trends in plumage colour and size are apparent. The blue wrens appear to
have originated as forest birds, probably in the New Guinea region, and to have evolved dull
female and male eclipse plumages for crypsis. It is suggested that the dull plumage on the
crowns of breeding males of assimilis may represent a similar trend.

INTRODUCTION

When the accumulated specimens of blue wrens, Malurus species, collected during
the five phases of the Harold Hall Australian Expedition were examined for com-
pletion of the final report, a re-examination was made of the forms within the chest-
nut-shouldered wren complex. The last revision had been that of Mack (1934).
Seven forms are recognized in the present study (Map i), these having been assigned
various taxonomic ranks during the past. In order to avoid any prior assumption
of taxonomic status a single specific or subspecific name has been used in referring
to each population in the following account. In addition to specimens collected
on the expedition, material already in the collection of the British Museum (Natural
History) and specimens borrowed from various Australian museums have been used.

COMMON CHARACTERS

All birds of both sexes have blue colouring, often rather dull, on the rectrices.
Males in breeding plumage have a black rump, a broad black collar posterior to the
ear-coverts and extending round the nape, and black lores. They also have scapular
patches of rather long chestnut-red feathers, the depth of colour on these tending to
vary with the intensity of the general plumage colour. The pale edges of the
tertials are also tinted chestnut-red. When not breeding males moult into an

Bull. Br. Mus. nat. Hist. (Zool.) 21, 8

C. J. O. HARRISON

eclipse plumage similar to that of females and immature birds of both sexes. I have
used the term "eclipse" plumage in preference to "non-breeding" plumage since
in some species of Malurus apparently adult males in breeding condition have the
plain plumage and are capable of breeding while in this plumage; thus while the
bright male colouring is undoubtedly a breeding plumage the converse is not always
true.

The plumage of females, immature birds, and eclipse males is plain brown or
blue-grey, according to the form. In those forms in which females have chestnut-
red on the lores and around the eyes, similar colour is present on immature birds of
both sexes. In these forms adult males do not regain this chestnut-red colour on
the lores after the first breeding plumage, although there may be some chestnut-red
on a few feathers immediately around the eye.

In moulting into the full breeding plumage, males of elegans and pulcherrimus
appear to acquire the full black lores while still otherwise in eclipse plumage, after
which bright blue feathers appear first immediately around the eye. The former
character was not apparent on a large series of specimens of assimilis and a few of
lamberti and rogersi, save for one brown-plumaged male specimen of assimilis with
completely black lores.

Except in amabilis, females and immature birds have reddish-brown bills. Males
in breeding condition have black bills, but may show some brown on a dark bill

rogersi
dulcis
amabilis
assimilis

lamberti

:::& pulcherrimus

066688$

';> elegans

MAP i . The ranges as indicated here are very tentative and should be regarded only as a
generalized diagram for the purpose of discussing relationship. The question marks
indicate regions where the ranges are in doubt, and within the areas shown populations
might be small and scattered.

THE CHESTNUT-SHOULDERED WREN 315

when in non-breeding plumage; while immature males approaching maturity show
an increasing spread of black.

Certain broad tendencies are apparent over the chestnut-shouldered wren group as
a whole. South-western birds tend to have purple breasts, north-eastern birds to
have white lores, northern birds to have blue-grey females and eclipse males, and
eastern birds to be bluer and less violet. Birds tend to become larger towards the
northerly and southerly limits of distribution.

FORMS OF THE CHESTNUT-SHOULDERED WRENS

Seven forms are recognized here and the accompanying table shows the major
differences in plumage between these. The forms are as follows.

Lores of ? Plumage of $ Flank of <$ Breast of <$

elegans chestnut-red rufous-brown and pale buff blue-black

greyish-brown

pulcherrimus chestnut-red olivaceus-brown pale buff dark violet

lamberti chestnut-red warm brown pale buff black

assimilis chestnut-red light brown pale buff black

rogersi chestnut-red light blue-grey white or greyish black

dulcis white light blue-grey white or greyish black

amabilis white dark blue very pale buff black

1. elegans. The Red- winged Wren.

This is restricted in distribution to the extreme south-west of Western Australia,
where it is found in thick cover associated with swamps, streams and lakes in areas
between Gingin and Warriup.

The male is the palest of these forms, being medium blue on crown and nape,
gradually changing on the sides of the head to very pale azure blue : paler still on the
ear-coverts where it has been described as "silvery". The back is an even paler
and more azure tint. The breast is blackish but with a strong violet tint producing
a blue-black colour; the latter most apparent when the bird is viewed frontally
with the bill raised and light falling directly on the underside, the breast appearing
deep violet with a black band along its lower edge. The hind-flanks and under
tail coverts are tinted with pale buff. The female is dark greyish-brown on the head,
and dark rufous-brown on the back and wings. Below it is light greyish-buff on
throat and breast and pale buff on belly and flanks. The lores are deep chestnut-red.

2. pulcherrimus. The Blue-breasted Wren.

This form appears in a zone north of that of elegans and mostly south of that of
assimilis. Its western limits are between the mouths of the Murchison and Namban
Rivers and its range extends through the mallee and wheatbelt in a south-easterly
zone to Warriup and Eucle, with an apparently isolate population on the Eyre
Peninsula.

The male is a deep violaceous blue on the head and deep violet on the back. The
forehead and sides of the head are more blue, becoming light blue on the ear-coverts.

316 C. J. O. HARRISON

The breast is glossy dark violet, brighter towards the edges of the sides, and with a
narrow black band along its lower edge. The flanks and under tail coverts are a
drab light buff. The female is dull olivaceous brown, with chestnut-red lores.
Below, the throat and breast are pale greyish-buff, the belly white, and the flanks
and under tail coverts light buff.

3. lamberti. Usually regarded as the eastern form of the Variegated Wren,
assimilis; but sometimes called Lambert's Wren.

This form occurs on the eastern seaboard, and the range appears to be the region
east of the Great Dividing Range, south to Sydney and north at least to the Brisbane
region, although there appears to be little evidence of what occurs near the coast
north of this. Further inland, specimens collected by Elsey (Macdonald and Colston
1965) on the Belyando River are certainly assimilis and the latter is said to have
been collected on the Dawson River (White 1916). These rivers, although inland
in the complex topography of the Dividing Range in mid-Queensland, drain towards
the east. Two female specimens collected at Bloomsbury, near Prosperine, on the
Wilkins Expedition (allowing for foxing and comparing them with material taken
elsewhere on the same expedition) are of the lamberti form, suggesting that the
latter extends well up the east coast.

Males of this form are deep blue, but not violet-blue, on the back and nape;
becoming paler blue on forehead, sides of head and ear-coverts, the last having, in
comparison with other light blue plumage, a slight azure tint. There are violet
tips to feathers bordering the sides of the black breast. Posterior flank feathers are
pale buff. Females, immature birds and males in eclipse are brown above, tinted
with warm buff on the rump; pale below with yellowish buff on flanks and belly;
and have deep chestnut-red on the lores and a narrow ring round the eye. The
brown colour on these birds is darker and warmer in tint than that of assimilis.

4. assimilis. The, Variegated Wren.

This form appears to occur from the Great Dividing Range and its ancillary ranges
in Queensland, westwards across the entire dry central region to the west coast,
north to the Gull of Carpentaria, the Roper River, and the Broome Area of Western
Australia, and south to the edges of the range of pulcherrimus but apparently not
as far as the south coast.

Males are violet-blue on the back, nape and crown, grading into deep blue on the
forehead and light blue, often slightly azure, on ear-coverts and around the eye.
Some individuals are less violet dorsally, lighter blue on the head, and more azure
on the ear-coverts. The posterior flanks are pale buff. There are violet tips to
feathers bordering the sides of the black breast. Of 33 adult males apparently in
full breeding plumage and not moulting, 17 showed an area of dull brownish colour
on the crown on the head, often resembling a distinct but irregular cap. Another
five show traces of this and only n have fully-coloured heads. Females, immature
birds and males in eclipse are a dull, light brown above, with a buffish rump.
Below they are pale buff, deepest on the flanks and almost absent from the throat
Adult males in this plumage, in addition to lacking the chestnut-red lores present on
the others, are much whiter below.

THE CHESTNUT-SHOULDERED WREN 317

5. rogersi. Usually regarded as the Western form of the Dulcet or Lavender-
flanked Wren, dulcis.

From specimens collected, and from others kindly lent by the Western Australian
Museum, this form occurs in the ranges of the Kimberleys from the Leopold Range
north to Napier Broome Bay and westward to the Ord River.

Dorsally males of this form are virtually indistinguishable from those of assimilis,
but do not show the extreme violaceous tint of some individuals of the latter and
tend to come about the middle of the range of variation of the blue and violet colours.
There is some individual variation. The wing feathers, both flights and coverts,
are a darker brown than those of assimilis and may show a faint bluish sheen.
This is apparent in fresh plumage, but an otherwise moulted male specimen showed
old wing feathers of a similar, sandier brown colour to those of assimilis, with just
a few darker new feathers. There are violet tips to feathers bordering the breast.
The belly and flanks are white, and of ten specimens only one shows a faint greyish
wash with even fainter violet tint which might have given rise to the vernacular
name. In the circumstances it would seem more correct to call this species "White-
flanked "rather than "Lavender-flanked" if it is desirable to refer to this aspect of the
plumage.

Females, immatures and males in eclipse plumage all have a dorsal plumage of
dull blue-grey, becoming paler blue on the sides of the head and neck. Males are
more blue and less grey, and very young birds are greyer, with a hint of brown, and
less blue. Wing-coverts are dark brown with greyish edges. Both females and
immature males show the chestnut-red lores that substantiate Mathew's rogersi.
Females show a very pale buff tint on the underside, while males in eclipse plumage
are almost white below.

6. dulcis. The typical Dulcet or White-flanked Wren.

This form is usually said to have a rather restricted distribution between the
Mary and King Rivers (the northern King River), in Northern Territory; (Storr
1967). Humphries (1947) refers to M. amabilis [=dulcis~\ being observed on one
occasion at Melville Bay, and more recently Rix (1970) has recorded this species at
Elsey Creek on the upper Roper River. It therefore seems possible that its true
range may be around, or through, the Arnhem Land Region.

Males appear almost indistinguishable from rogersi but in fresh plumage appear to
show a more obvious bluish wash on the wing feathers, particularly the coverts,
making these appear darker in colour. Females and immature males differ from
rogersi in being slightly darker dorsally, but more conspicuously in having the lores
and a narrow ring round the eye creamy-white and not chestnut-red. Ventrally
the very pale buff colour is apparent but is combined with a faint grey tint to give
the plumage of the underside a rather drab appearance.

7. amabilis. The Lovely Wren.

This is another form with apparently limited distribution, occupying habitats
bordering the rainforest along the north-east Queensland coast between Cape York
and Cardwell, but it has also been recorded (Thompson 1935) from the Gulf coastal

318 C. J. O. HARRISON

regions on the Lower Edward River where it was noted and collected in "dense dry
scrubby country on raised beaches".

Males are most similar to lamberti, but are a lighter, clearer blue on the back and
head, without any definite violet or azure tints, the ear-coverts being similar in
colour to the rest of the head. The violet tips to feathers at the sides of the breast are
restricted to a few vestigial tips. The flanks show some pale buff. The pale edges
to wing feathers show a more distinct blue wash, and these feathers are otherwise
very dark in colour, in some instances practically black, and in the case of the
lesser coverts usually black.

Females, immature males and males in eclipse resemble those of dulcis in having
creamy-white lores and eye-rims, but differ in the much darker colour of the dorsal
plumage. The latter is a dark and rather dull blue, becoming a little lighter on the
forehead, and with conspicuous light blue ear-coverts. Both females and males in
eclipse show black bases to the feathers across the upper mantle which are wholly
black in the breeding males. The bill is black in both sexes. Wing feathers are
dark with a dark blue wash. Below the plumage is very pale buff, whiter on the
throat, and whiter overall on males. The juvenile is browner and less blue on the
back, and has a dark, blackish-brown bill.

VARIATION WITHIN FORMS

Individual variation is apparent within assimilis, rogersi and amabilis. It is
assimilis, with its very extensive range, which shows the greatest variation. Indi-
viduals showing some variation were described as new races and originally nine were
recognized within the range of this form, but these were reduced by Mack (1934) to
three (Map 2). On the basis of the material examined, I cannot agree that there
are three readily recognizable subspecies within this area, but there is some tendency
toward broad trends in colour variation within the very large range occupied by
this form.

As already mentioned, the only obvious difference is for some individuals to have
paler heads. In comparing the blue colour it is necessary to have specimens side
by side at the same angle to the light, since the incidence of light can affect the
apparent hue of structural colours of feathers.

The original description of assimilis was based on one of the darker south-eastern
specimens. Grant (1909) described bernieri, from Bernier Island off the west coast,
differing in that the ear coverts of the male were a dark blue. In a specimen examined
they are undoubtedly dark, but match well with those of a specimen from near
Lake Frome in South Australia, and one from near Hughenden in northern
Queensland. The third race, mastersi was described by Mathews (1912) from a
specimen from Alexandra, Northern Territory, with more azure blue ear-coverts.

If a dark specimen is compared with a light one the difference is quite obvious,
but it was found possible to lay out a series of adult males which showed a consistent
gradation of forehead and ear-covert colour, from one extreme to the other, with
no division at any point to suggest that more than one population was involved and
there was no clear geographical pattern. The specimens, beginning with the deepest

THE CHESTNUT-SHOULDERED WREN 319

colour, were from the following localities: i, South Australia (unlocalized older
specimen, probably from south-east settlement); 2, Bernier Island, W.A. ; 3, 30
miles west of Lake Frome, S.A. ; 4, near Hughenden, Qld; 5, 15 miles west of Lake
Frome, S.A. ; 6, Bourke, N.S.W ; 7, Finke River, N.T. ; 8, Tambrey, W.A. ; 9, Point
Cloates, W.A. ; 10, Prairie, Qld; n, Little Desert, Vic,; 12. Warburton Mission,
W.A. ; 13, Tambrey, W.A.; 14, Upper Gascoyne, W.A. ; 15, Moonlight Creek, Qld.;
16, Upper Gascoyne, W.A.; 17, Fitzroy River, W.A. ; 18, Upper Gascoyne, W.A.
(see map 2).

If one were to follow Mack (1934) nos. 6 and n would form his dark subspecies;
no. 2 another; nos. i, 3, 5, 7-9. 12-18 would form a pale subspecies; and nos. 4 and
10 would be part of an indeterminate population. Mack himself appeared uncertain
of the precise limits of the forms which he recognized and in his map of subspecific
distribution (map 2) did not show precise areas but indicated lobes from a main
mass. Condon (1951), in discussing South Australian birds, abandoned head and
back colour as diagnostic characters and disagreed with Mack's views on subspecific
limits. Ford (1966) noted the variability of the head colouring in Western Australian
birds.

There is some evidence of a general tendency for birds with deeper and darker
blue colour to occur on the eastern and extreme western edges of the overall distri-
bution, and for paler birds to be most frequent in the desert areas of Western Aus-
tralia and towards the north-western limits of distribution. Perhaps Gloger's rule

MAP 2. The dotted line indicates Mack's version of the distribution of races of M. lamberti :
A, M. I. lamberti; B, M. I. assimilis; c, M. I. master si; D, M. I. bernieri. The figures are
those indicating individuals in the section on variation within forms. The black dots
indicate the localities of variant individuals.

320 C. J. O. HARRISON

is involved. This clinical trend does not, however, appear to allow convenient
subdivision and, in addition, there is evidence of variability between individuals in
small areas as well as over a wide range. Nos. 8 and 13 from the above list were
collected on the same occasion from the same party of birds, and there are similar
variations within specimens from single localities at Fitzroy River, Western Australia,
and in northern Queensland. T. Carter (Mathews 1922-3) noted a bird taken on the
Minilya River as "blue" on the back, although most individuals from the west
coast area were distinctly violet.

Other characters for separation have been suggested. Condon (1951) stated that
master si could be distinguished by its clear white abdomen. This is difficult to
determine in the skins of very small birds prepared by a variety of hands, but it
was not apparent in the material now examined (46 males) and where some slight
difference was apparent it did not relate to the earlier subspecific divisions.

Condon and others also refer to differences in the relative lightness of the brown
colour of the wings. The glossy black and blue body plumage appears to be stable
but the brown plumage of the wings shows a definite tendency to fade on the living
bird and most of the variation apparent in specimens examined due to the differing
age of the feathers, the fresh new feathers being darkest. In addition there is some
change due to foxing in older specimens. The wing moult and body moult do not
appear to be closely synchronised, and birds which show complete breeding plumage
on the body may still be actively moulting and growing wing feathers. The relative
depth of colour on the wings does not therefore provide a consistent and useful
taxonomic character.

The difference in plumage between individuals within assimilis are so slight and
gradual that there seems no reason to suppose that more than one unit is involved
with some local differentiation beginning to occur. I am of the opinion that this
should be treated as a single form, and assimilis North, 1901, is the oldest name;
mastersi Mathews, 1919, and'bernieri Grant, 1909, being synonyms.

MAJOR VARIATION IN THE RED-SHOULDERED WREN GROUP

Among specimens of rogersi a male from Kulumburu (W.A. Museum, A 8884)
was a much lighter blue on the head, and similar in this respect to a specimen of
assimilis (B.M. no. 1964. 60.585) from Moonlight Creek, north Queensland.

There is some variation in the blue tint of the head of breeding males of amabilis.
Mack (1934) separated birds from the southern half of the range as a subspecies,
clams, on the grounds that they showed lighter blue colouring. An examination
of specimens, including some lent by the Queensland Museum, revealed a small
difference between extremes. Birds recently collected from Tully were a lighter
blue than some early specimens from Somerset and Port Albany, and the latter showed
a slight violaceous tint to the deeper blue which was a little more distinct on some
northern birds from the Queensland Museum. Recent specimens from Ayton,
towards the middle of the range of this form, appeared to be intermediate in character.
The total difference appears to be small and clinal in character and on the material
at present available I would regard clams Mack, as a synonym of amabilis.

THE CHESTNUT-SHOULDERED WREN 321

VARIANT INDIVIDUALS

In addition to the variations already described there are some variant specimens
which, from their appearance and locality, (map 2) may throw some light on the
relationship between various forms.

The most conspicuous of these is an adult female assimilis (B.M. no 1964.60.578)
collected on the Norman River south of Normanton, Queensland, at the south-
east corner of the Gulf of Carpentaria. This bird shows a pale greyish tint over the
whole body plumage, particularly noticeable on the sides of the head and neck.
The lores are the normal chestnut-red colours and the retrices dull blue. There is
no suggestion that this variation is due to wear or fading. Normal brown plumage
contains both eumelanin and phaeomalenin and these are not differentially affected
by exposure to light, (Harrison 1963). The bird was collected from a party of
eight individuals and another female and an immature male collected at the same
time appear to be normal.

Two other females of assimilis (B.M. nos 1969.4.415, 417) show similar, although
less marked, tendencies towards greyish plumage. These are the only two females
taken on the Roper River, Northern Territory (14 I5'S., 135 3' E.). On these
the grey tint is superimposed on the normal brown plumage to give a colder, greyer
tint, but the sides of the head and neck are noticeably grey, and the rump has an
olive tint.

A minor plumage variation found during close examination was the presence of a
few dark lesser covert feathers on some males of assimilis, the coverts of this form
normally being brown. Occasional blackish covert feathers occurred on a male
(B.M. no. 1964.60.674) from Norman River, at the same "locality as the female
described above, but from a different party of birds; and on a male (B.M. no 1964.
60.585) from the south of Moonlight Creek, on the south-east of the Gulf of Carpen-
taria. Males showing some blue tips to feathers occurred at Moonlight Creek
(B.M. no 1964.60.581), Roper River (1969.4.414, 416), and at Mount Anderson
(1964.4.443, 450) in the south-western Kimberleys.

These variants may be significant in view of their distribution. The greyish
females of assimilis occur on the edge of its range, in one case near amaUlis and in
the other near dulcis, both of which have blue-grey females. The variant wing-
coverts similarly occur on individuals on the northern edge of the range, birds with
blackish coverts at Norman River and Moonlight Creek being near amabilis which has
similar coverts ; and birds with blue covert tips at Roper River and Mount Anderson
being near to dulcis and rogersi respectively, both of these having blue on the converts.
The exception is the second male from Moonlight Creek which has the few covert
tips blue and not blackish.

ZOOGEOGRAPHICAL DISTRIBUTION PATTERN

The type of distribution shown by the chestnut-shouldered wren complex appears
to provide a good example of the refuge concept suggested by Keast (1961) and to
show the utilisation of a greater number of refuges than do the other superspecies
groups which he instances. Keast did use these birds as an example but used the
races as mapped (map 2) by Mack (1934) and hence found the parallels with other

322 C. J. O. HARRISON

species groups less obvious than they should have been. There is, in fact, a ring of
forms, most of which still occupy limited peripheral areas (map i). On the eastern
edge of Australia is lamberti, amabilis is in the north-eastern peninsula, dulcis in the
north of Northern Territory, rogersi in the Kimberleys of the north-west, and
elegans in the south-west corner. Since pulcherrimus is present on the Eyre Penin-
sula as well as in the south-west it is more likely that, rather than having evolved
in competition with elegans, it differentiated in or somewhere near the former region
in a refuge demanding tolerance of drier conditions and higher temperatures than
did the south-west refuge; and therefore, when conditions ameliorated, it would
have been able to spread westwards into similar habitats bordering the range of
elegans.

The last form, assimilis, presents a slight problem. There are several potential
refuges which might have been available (Keast, 1961) and are not occupied by
other members of this group, but from its present distribution it is reasonable to
assume that the differentiation would have occurred in a refuge where conditions
were both drier and warmer than in some others. The extreme south-east is there-
fore unlikely since conditions there are likely to have been both moister and cooler.
The two remaining likely areas are the Hamersley region of coastal Western Australia
and the central ranges. The differentiation in isolation apparent in other taxa
(Keast 1961) occurs less frequently, in the latter area and the Hamersley region
would seem the more likely of the two. The adaptation to such a refuge would
give this form the slight advantage that would enable it to exploit more rapidly and
successfully the gradual amelioration of extremely arid conditions of the central
area and to spread to produce the apparent "Eyrean" distribution (Spencer 1898).
Keast (1958) has pointed out that many forms with this distribution are derived
from western isolates of species with more extensive distribution at an earlier period.

HABITAT PREFERENCE

From the limited distribution of some of these forms, they may have evolved
some degree of habitat preference which might limit any subsequent spread. In the
south-west elegans has a limited distribution in low cover bordering fresh water
swamps or streams; while pulcherrimus is a species of sandplain scrub and mallee.
Of the northern forms, both rogersi and dulcis occur in regions of ranges and plateaus
and appear to be mainly confined to places where the floors and sides of sandstone
or granite gorges have low scrubby vegetation and where natural breaks or dis-
continuities in the rock of the ranges are accompanied by vegetation including low
bushes and spinifex. The other extreme is shown by amabilis which occurs on the
outer edge of rain-forest or in suitable low cover in open forest adjoining it. The
information on lamberti is poor but it appears to occupy thick, shrubby growth in
fairly moist habitats including the thick shrub layer of forest, and in this respect
would seem to show some similarity to amabilis.

As might be expected from its considerable range, assimilis shows a wider habitat
tolerance. In general it tends to be a bird of shrubby growth bordering water-
courses in drier regions. It may, however, move into the sparse vegetation of
sandstone ridges, occupying a similar habitat to rogersi and dulcis in similar regions,

THE CHESTNUT-SHOULDERED WREN 323

and although from their field observations B. M. Booth and D. Freeman are of the
opinion that in such places it is less likely to venture onto bare rock than are the last
two forms this would not prevent it from sharing the same cover were it to occur
with them. At the other extreme it extends into the shrub growth along rivers and
creeks in open forest; and down into thickets, riverine forest and the edges of
mangroves.

In Western Australia near Carnarvon, where this form occurs in the scrub on
dunes among saltbush, pairs were seen out in the mangroves apparently disputing
territory over rising seawater. The mangroves in this area form a narrow belt
separated from the beach by a tidal lagoon up to about half a mile wide in places.
Within the mangroves, which are low and open on the landward side, becoming
taller and thicker to seawards, there are small crescentic beaches with a sparse
growth of herbaceous plants. Although it seems likely that the presence of pairs
in such a place may have been due to a lack of suitable territories due to overpopu-
lation, and the fact that these were pairs rather than parties suggests that they were
young or breakaway units searching for new areas, their presence nevertheless indi-
cates the readiness of this form to attempt to occupy a wide range of habitats.

This wide habitat tolerance in assimilis does suggest that the ecological require-
ments of the birds may be relatively simple and that, at least where the northern forms
are concerned, the apparently narrower habitat preference of various other forms may
be simply due to the fact that they are occupying the ecological equivalent within a
more specialised and less varied biotope. Certainly the habitat tolerance of assimilis
is such that were it to come into contact with other forms it seems likely that it
would occupy the same niche, and the apparent minor variation in habitat would be
unlikely to act as a barrier between them.

THE INTERACTIONS AND TAXONOMIC STATUS OF THE FORMS

Where a number of similar allopatric forms exist it is always difficult to determine
their precise taxonomic relationship. In the present instance the recognition of
five species for the seven forms appears to be an accidental result of the successive
description of the forms and their similarity to those already known at the time of
their discovery.

The situation in the south-west has been well investigated (Serventy 1951, Ford
1966, 1969). Here elegans and pulcherrimus have contiguous ranges and pulcheri-
rimus and assimilis are sympatric in the north-west part of the former's range.
Ford (1966) has evidence that these pairs encounter each other in the field; but
elegans and assimilis do not meet. In such encounters these three forms appear to
ignore each other and behave as good species. Where pulcherrimus and assimilis
are sympatric they occur in the same biotope and show a mosaic distribution, but
it is not certain whether this indicates interspecific intolerance or differing responses
to microhabitats (Ford 1966).

The plumage of breeding males appears adequate to ensure specific recognition,
the combination of colour on breast, head and back being conspicuously different in
the three (table i). A combination of pale azure and blue-black is present on
elegans; violet-blue and dark violet on pulcherrimus; and lighter violet-blue and

324 C. J. O. HARRISON

black on assimilis. These colours would be particularly conspicuous in any frontal
displays. The interspecific variation in colour of females, young and eclipse males is
relatively slight, but this might aid specific recognition. There may be other
differences. Ford (1966) refers to a detectable difference in the voices of pulcherrimus
and assimilis. Size differences are relatively slight, with a clinal increase in a
south-westerly direction; and I doubt if the differences in bill-size given by Ford
would be sufficient to produce the variation in food selection which he suggests.

The situation is more complicated in the northern half of Australia where there
are five recognizable forms lamberti, assimilis, rogersi, dulcis and amabilis pre-
viously recognized as three species lamberti I assimilis, roger si j dulcis and amabilis.
These five forms show slight size variation of a clinal type, the more northerly
being a little larger. Since within the whole chestnut-shouldered wren group
size tends to increase towards the northern edge of the distributional range as well
as the southern edge it seems inadvisable to suggest the latter as an example of
Bergmann's rule.

Apart from this, the characters which differ between forms and could be used for
recognition are those of plumage colour. In Malurus species generally specific and
subspecific variations are usually most apparent in the breeding plumage of the
male. In the present group although the dorsal blue colour varies, the range of
variation within assimilis encompasses that of both rogersi and dulcis. In sequence
from blue to violet-blue the arrangement would be amabilis lamberti assimilis
(inc. rogersi and dulcis) ; the difference between the last two of the three, usually
regarded as conspecific, being greater than that between assimilis and the forms in
parentheses which are regarded as good species.

The only other obviously variable plumage character of breeding males, which
has been used for separation of forms, is the flank colour (table i). The posterior
flanks show a variable amount of pale buff on lamberti and assimilis, slightly buff
colouring on amabilis, and are usually white, with a purplish-grey wash on some
individuals of rogersi and dulcis. One or two specimens of the last form show a
small amount of pale buff, particularly around the upper thighs, but since buff is
present on females and eclipse or immature males, this may only indicate incomplete
assumption of full male plumage.

The female plumage shows greater distinctiveness in this group, varying between
brown and blue-grey ; lamberti being warm brown, assimilis normally a paler, duller
brown, rogersi and dulcis pale blue-grey, and amablis deep blue-grey. The apparent
clear-cut distinction is blurred a little by the existence of the greyer variant individ-
uals of assimilis. If these are included in the sequence there is a much smoother
gradation and the most relevant differences would appear to be the darker colour of
amabilis and the change from chestnut-red lores on lamberti, assimilis and rogersi to
white lores on dulcis and amabilis.

The mainly allopatric distribution makes it difficult to judge the amount of
interaction that might potentially occur between these northerly forms. Mack (1934)
suggested that assimilis and lamberti interbred in northern Queensland, but from
his account it seems possible that he did not make allowance for variation within
assimilis, and a more satisfactory investigation of this is still needed. At the oppos-

THE CHESTNUT-SHOULDERED WREN 325

ite extreme of the range of the latter the fifth phase of the Harold Hall Australian
Expedition found both assimilis and rogersi near Mount Bell in the Leopold Range,
one of the more level plain and the other in the gorges of the range, but in types of
habitat in which they might well encounter each other. Intergrades between the
two would be very difficult to identify. Assuming that interbreeding produced
individuals with intermediate plumage colour, this would only involve the buff
on the flank of the male and the body colouring of birds other than breeding males.
Any male bird with buff on the flanks would probably pass as assimilis. Inter-
mediates when female or immature should be recognizable and would probably
resemble the greyer variant individuals of assimilis already described. If these
single plumage characters were controlled by a simple pair of dominant and recessive
alleles recognition of a hybrid might be impossible.

Rix (1970) has recently recorded a party of assimilis in a clump of teatree on
Elsey Creek with a party of dulcis (referred to by Rix as amabilis) in an area of tall
grass with a few small shrubs only about twenty yards away. There is therefore
a potential zone of interaction between assimilis and dulcis also; but theoretically
none between assimilis and amabilis (map i.) However, the presence of variant
individuals of assimilis apparently showing the plumage character of the other
two to some degree, at points in its range nearest to the known ranges of the
other two forms, suggests either that the factors responsible for these plumages
are environmental ones which may act upon individuals of assimilis in the same
regions and tend to select for similar characters (although in such circumstances one
might expect all individuals in an area to show some evidence of this) or that there
is some interbreeding within the areas where the forms approach each other. If the
latter is true then it would suggest that the distribution of amabilis is more extensive
than our present knowledge indicates. To suggest such interaction presupposes that
there are not fully effective barriers of ecology or species-specific recognition between
the forms. The obvious ecological barrier of habitat preference has already been
discussed and it is suggested that the wide habitat tolerance of assimilis could
potentially bring it into contact with the other forms. Recognition would be based
on signals of behaviour, voice or plumage pattern. Such little evidence as we have
suggests that these various forms are similar in their behaviour and that their
voices are indistinguishable, although in view of Ford's (1966) comment on the voices
of two south-western forms a more critical appraisal of this character might be
helpful.

In the south-western forms, where plumage colour of the breeding males appears
to be a good specific character, adjacent forms show marked variation in the com-
bined colouring of head, back and breast. This suggests that such signal colouring
is associated with frontal displays, and such postures have been described (Rowley
1964) in the Superb Blue Wren, M. cyaneus, the only adequately studied species.
The present forms all have black breasts, with slight variation in the extent of the
violet edge at either side. The head and back colouring is indistinguishable and
variable in three forms and it is therefore difficult to argue that the slight differences
in the shade of blue shown by the other two is of any great significance. The
flank colour used to separate forms would not appear to be used in displays and,

326 C. J. O. HARRISON

in some at least, might be actually concealed by the wings. There would therefore
seem to be no reason to regard the male breeding plumage as obviously specifically
isolating.

An alternative isolating factor would be the recognition by the male of differences
in the female colour, the grey or brown plumage, the presence or absence of chestnut-
red lores, or the bill colour. Heterogynism variation in females but not in males
occurs in races of some species but is comparatively rare. Mayr (1942) records it in
three races of Pachycephala pectoralis in the Solomons, but since these are on different
islands the question of recognition or interaction would presumably not arise.
Hellmayr (1929) and Zimmer (1931 et seq.) also found it in races of some species of
Formicariidae in South America, but the implication of this in relation to recognition
does not appear to have been examined. There is therefore no useful information
on the use of heterogynism in intraspecific recognition. Encounters of forms in
which chestnut-red lores are present with those which lack them might produce some
confusion since in the forms that possess them the absence of these usually indicates
that a bird in brown or blue-grey plumage is an adult male in eclipse. Similarly
a black bill on a bird in eclipse plumage usually indicates a male, but females of
amabilis also have a black bill.

In summary, there are two very distinct forms of the chestnut-shouldered wrens
in the south-west which can be regarded as separate species; but the inter-relation-
ships of the remaining five forms are obscure. Each has a slightly different com-
bination of plumage characters. The selection of particular characters for the pre-
vious separation of species seems to have been arbitrary, and each form would appear
to represent a separate isolated evolutionary unit. The more recent taxonomic
treatment of the peripheral rings of isolate forms, of which there are many in the
different taxa on the Australian sub-continent, has been to regard these as species,
except where secondary re-integration occurs in forms which spread and re-encounter
each other, in which case they are regarded as subspecies. A typical example of
the latter is Keast's (1961) interpretation of relationship in the Australian forms
of Sittella.

In the case of the chestnut-shouldered wrens there are five distinct forms of
equal standing, all of which could be regarded as specific or near-specific entities.
It is possible however that speciation has been incomplete and that they might
interbreed and re-integrate freely if they came together again. The extensive
spread of assimilis has increased the likelihood that this might occur, and the
existence of variant individuals suggests that it may be taking place. If subsequent
field investigation shows that this is so, and if more detailed study of these forms
confirms that there are no barriers, of the type discussed, to prevent this occurring,
then it might be preferrable to regard them as subspecies within a single species.

APPARENT EVOLUTIONARY TRENDS

The genus Malurus appears to have affinities with a group of genera in New
Guinea (Harrison and Parker 1965, Harrison J_969a). The deep blues, black, and
chestnut-red of these birds is a colour combination which appears more usually to
originate in moist tropical forest; and from this, and from what we know of the

THE CHESTNUT-SHOULDERED WREN 327

general zoogeographical trends within this region, it seems probable that Malurus
as a unit may have invaded Australia from this direction and subsequently evolved
and radiated within the Australian environment. In the New Guinea group as a
whole females are similar in plumage colour to males, or as richly coloured. The
production of the rather drab female plumage, and of an eclipse plumage in the non-
breeding males appears to have occurred in Australia, and it would seem most
likely that this is an adaptation for crypsis in a generally more open environment
where the more brightly coloured bird is vulnerable to predation. In the case of
the Blue-and- White Wren, M. leucopterus, there appears to have been a suppression
of bright male plumage in subordinate males (Harrison 1969^, the alternative
possibility being that of very high predation on males in breeding plumage. The
species is one that occurs in arid areas with relatively sparse cover, and the blue and
white male is very conspicuous.

In view of this apparent tendency to reduce the conspicuous colour of the males of
this taxon in the more open areas, retention of the dull eclipse plumage on the crowns
of so many of the collected male specimens of assimilis may be significant. With
such a bird in its normal posture with tail cocked acutely and head back a little,
the back is relatively less conspicuous and to an aerial avian predator the crown
of the head must be the most obvious mark. A dull crown might thus carry a strong
selective advantage.

ACKNOWLEDGEMENTS

I am very grateful for the loan of material to the Western Australian Museum,
the Arid Zone Research Institute of Northern Territory, the C.S.I.R.O. Division of
Wildlife Research at Canberra, and the Queensland Museum. I should also like
to acknowledge the help and information that I have received from Major B. M.
Booth, Mr. D. J. Freeman, Mr. C. Frith, Mr. I. C. J. Galbraith, Mr. S. A. Parker,
Dr. D. W. Snow and Dr. G. M. Storr.

REFERENCES

CONDON, H. T. 1951. Notes on the birds of South Australia; Occurrence, distribution and

taxonomy. S. Aust. Orn. 20 : 26-28.
FORD, J. 1966. Taxonomy and variation of the Chestnut-shouldered wrens of Western

Australia. Emu 66 : 47-57.

1969. Range limits of the Blue-breasted Wren in Western Australia. Emu 68 : 283-284.
HARRISON, C. J. O. 1963. Grey and fawn variant plumages. Bird Study 10 : 219-233.
ig69a. The affinities of the Blue Wren genus Malurus and related genera; with special

reference to the grass-wren genus Amytornis. Emu 69 : 1-8.

i96gb. Helpers at the nest in Australian passerine birds. Emu 69 : 30-40.
HARRISON, C. J. O. & PARKER, S. A. 1965. The behavioural affinities of the Blue Wrens of

the genus Malurus. Emu 65 : 103-113.
HELLMAYR, C. E. 1929. On heterogynism in Formicarian birds. /. Orn. Lpz., Ergbd.

2 : 41-70.
HUMPHRIES, C. P. 1947. Among the birds at Melville Bay. Emu 47 : 130-136.

328 C. J. O. HARRISON

KEAST, A. 1958. The genus Psophodes Vigors and Horsfield, and its significance in demon-
strating a possible pathway for the origin of Eyrean species from Bassian ones. Emu
58 : 247-255.

1961. Bird speciation on the Australian continent. Bull. Mus. Comp. Zool. Harvard

123 : 303-495-

MACDONALD, J. D. & COLSTON, P. R. 1965. J. R. Elsey and his bird observations on Gregory's
overland expedition, Australia, 1856. Emu 65 : 255-278.

MACK, G. 1934. A revision of the genus Malurus. Mem. Nat. Mus. Viet. 8 : 100-125.

MATHEWS, G. M. 1922-23. Birds of Australia. Vol. 10. Witherby; London.

MAYR, E. 1942. Systematics and the origin of species. Columbia U.P. : New York.

OGILVIE-GRANT, W. R. 1909. [Three new species of birds from West Australia]. Bull. Brit.
Orn. Cl. 23 : 72.

Rix, C. E. 1970. Birds of the Northern Territory. South Australian Ornithologist 25 : 147-
190.

ROWLEY, I. 1964. The life history of the Superb Blue Wren, Malurus cyaneus. Emu
64 : 251-297.

SERVENTY, D. L. 1951. The evolution of the Chestnut-shouldered Wrens (Malurus}. Emu
51 : 113-120.

SPENCER, B. 1898. Report of the work of the Horn scientific expedition to Central Australia.
Narrative. Dulau: London.

STORR, G. M. 1967. List of Northern Territory Birds. Spec. Publ. W. Austr. Mus. No. 4.

WHITE, H. L. 1916. A new Wren-Warbler, Malurus lamberti dawsonianus (Barnard's Wren).
Emu 16 : 69.

ZIMMER, J. T. 1931 et seq. Notes on Peruvian birds. Amer. Mus. Novit. 500 et seq.

Dr. C. J. O. HARRISON

Department of Zoology

BRITISH MUSEUM (NATURAL HISTORY)

CROMWELL ROAD

LONDON, S.W.7

A NEW SPECIES OF LIRONECA (ISOPODA;

CYMOTHOIDAE) PARASITIC ON CICHLID

FISHES IN LAKE TANGANYIKA

By R. J. LINCOLN

INTRODUCTION

ONLY three species of parasites belonging to the family Cymothoidae have been
recorded from freshwater localities on the African continent. The first of these to
enter the literature was Ichthyoxenus expansus, described by Van Name in 1920 as a
parasite on the gills of the citharinid fish Eugnathichthys eetweldii Boulenger. This
isopod has since been widely recorded from the waters of the Congo river basin
(Monod, 1931 ; Dartevelle, 1939; Brian & Dartevelle, 1949 and Grosse, 1963). There
has been much discussion in the past about the validity of the genus Ichthyoxenus
to which expansus was referred. Miers (1880) expressed the opinion that the two
genera Ichthyoxenus and Lironeca were indistinguishable, except that the former
was exclusively freshwater. Van Name (1920) considered the wide separation of
the localities from which the species of Ichthyoxenus had been described as evidence
that they were not a monophyletic group, but were instead convergent forms. In
this view the freshwater habitat has been adopted independently a number of times
by some ancestral marine genus such as Lironeca. Monod (1931) also mentions the
above points, but despite this the name Ichthyoxenus still seems to survive for
certain freshwater species. More recently, Fryer (1965) adopts the view that the
freshwater species have been derived from some common marine ancestor, and that
morphologically they are indistinguishable from the widespread marine genus
Lironeca. Fryer thus refers expansus to Lironeca, and considers Ichthyoxenus of
Herklots 1870 to be a junior synonym of Lironeca Leach 1818.

Two further species are described by Fryer from Lake Tanganyika, Lironeca
tanganyikae Fryer 1965 and Lironeca enigmatica Fryer 1968. The former species,
tanganyikae, has been recorded only from the mouth cavity of the littoral cichlid
fish Simochromis diagramma (Giinther), while enigmatica was taken from the body
and gill cavity of the clupeids Limnothrissa miodon (Boulenger) and Stolonothrissa
tanganicae Regan.

An examination of a collection of the cichlid fishes Lamprologus elongatus
Boulenger and Lamprologus pleurostigma Boulenger from Lake Tanganyika has
provided a number of isopod parasites, with both mature male and female stages
present. These specimens represent a hitherto undescribed species which I name
Lironeca africana.

Bull. Br. Mus. nat. Hist. (Zool.) 21, 8

330 R. J. LINCOLN

Lironeca africana sp. nov.
(Text-figs la-g, 2a-j, 3a-q)

MATERIAL EXAMINED. Holotype female, length u-o mm, ovigerous: allotype
male found in association with holotype, length 5-0 mm: paratypes, 5, $$, length
7 -0-14 -o mm, width 3 -5-5 -o mm ; 4, <$$, length 4 -0-5 -5 mm, width i -0-2 -o mm. All
specimens deposited in the British Museum (Natural History), accession numbers;
holotype, 1970 : 438; allotype, 1970 : 439; paratypes, 1970 : 440.

FEMALE, (holotype). Body size, length n-o mm, width 4-5 mm, height including
the marsupium 4-0 mm. Shape of body in dorsal, ventral and lateral view given in
Figs la, 2a, ib respectively. Head, ratio of length to maximum width about
i : 1-5, shape triangular, broadly rounded at the front and curved slightly down-
wards towards the base of the antennae ; eyes large, laterally placed ; head somewhat
sunken into the peraeon ; chromatophores evenly distributed on dorsal side of head.
Peraeon tergite i about i| times the length of tergite 2, tergites 2-6 subequal,
tergite 7 only half length of tergite 6; peraeon tergite i, antero-lateral projection
reaching forward to mid-eye level, anterior and posterior margins broadly rounded ;
peraeon tergite i almost flat dorsally, tergites 2-7 becoming progressively more
convex ; maximum width of peraeon is reached by coxal plates of segment 4 ; coxal
plates reach progressively nearer to posterior margin of tergites from 1-7 ; posterior
margin of peraeon tergite 7 broadly rounded. Pleon immersed a little in peraeon
tergite 7, all segments free laterally, length of segments increasing slightly from 1-5 ; pos-
terior margin of pleon segment 4, and especially 5, sinuous. Telson, length twice width,
broadly rounded, with ridge running from antero-lateral corners towards the centre.

Appendages; antenna i and 2 each with 7 segments, slender, rounded and only
just reaching beyond forward projection of peraeon tergite i, antenna i slightly
longer than antenna 2 (Fig. 2b, c) ; mandible (Fig. 2d) with slender inscisor process
pointed at tip, molar process rounded, palp 3-segmented with 2-3 apical setae;
maxilla i (Fig. 2e) slender, carrying 4 recurved spines on distal tip ; maxilla 2 (Fig. 2f)
small, bilobed, each lobe bearing a pair of hooked spines; maxilliped (Fig. 2g) well
developed, broad, with a 2 segmented palp terminating in 4 strongly recurved spines.
Peraeopods 1-7 of similar shape (Fig. ic, d), basis and ischium strongly developed in
anterior peraeopods, less so in posterior peraeopods, merus and carpus very short,
propodus prominent, dactylus strongly curved and pointed, apex of dactylus fitting
into a carpal groove, very well formed articulation between dactylus and propodus,
no spines present on the peraeopods. Pleopods fleshy, leaf-like, without marginal
setae (Fig. ic, f) ; uropods (Fig. ig), inner ramus just a little longer than outer, both
rami apically rounded, projecting fractionally beyond margin of telson.

MALE, (allotype) found associated with holotype. The male is very much smaller
than the female, length 5 mm, width 2 mm ; general form of the body and appendages
resemble the female, and reference will therefore be made only to the main points of
difference. Shape of body in dorsal view as in Fig. 2h. Head with rounded eye
lobes, produced forward into a short, somewhat square-ended rostrum, only weakly
immersed into peraeon; peraeon slightly asymmetrical, tergite i is i| times length

LIRONECA (ISOPODA; CYMOTHOIDAE)

FIG. i. Lironeca africana sp. nov. $ Holotype: a, body entire, dorsal view; b, body entire,
lateral; c, peraeopod i left, ventral view; d, peraeopod 7 left, ventral view; e, pleopod i
left, ventral view; f, pleopod 2 left, ventral view; g, uropod left, ventral view. Bar scale
i mm.

332

R. J. LINCOLN

FIG. 2. Lironeca africana sp. nov. a-g $ Holotype: a, body entire, ventral; b, antenna i ;
c, antenna 2; d, mandible, e, maxilla i; f, maxilla 2; g, maxilliped; h-j <$ allotype:
h, body entire, dorsal; i, pleopod i, left; j, pleopod 2, left. Bar scale i mm.

LIRONECA (ISOPODA; CYMOTHOIDAE) 333

of tergites 2-7, maximum width at the level of segment 3, posterior margin of tergite
7 deeply rounded ; paired penis lobes present on ventral surface of peraeon segment 7 ;
chromatophores on both dorsal and ventral surface of peraeon; body less convex
than the female. Pleon segment i immersed laterally in peraeon, segments 2-5
free; telson broad and flat, length about half of width, posterior corners round
becoming straight in the centre.

Appendages; antenna i (Fig. 2h) rounded in cross section, 7 segmented; antenna
2 much narrower and consisting of 8 segments; mandible, maxilla i, maxilla 2 and
maxilliped as in female except the apical hooks are much less pronounced; pleopod
2, endopod, bearing appendix masculina (Fig. 2J), pleopod i, fleshy and leaf -like
(Fig. 2i).

First marsupial stage. This is mentioned only briefly as it is the structure of the
second marsupial stage which is of greater interest. Body shape as in Fig. 3m;
head not immersed in peraeon; peraeon tergite i with markedly sinuous posterior
margin, segment 7 reduced in size and continuous with pleon; telson long and
rounded, no marginal setae.

Appendages; antenna i consists of 7 segments, much dilated, without aesthetascs;
antenna 2 of 8 segments, slender, slightly longer than antenna i. Peraeopods 1-6
of similar size (Fig. 3n, o), no hooks or spines present. Pleopods 1-5 fleshy, no
marginal setae; uropods projecting well beyond extremity of telson, no marginal
setae or terminal spine (Fig. 3p, q).

Second marsupial stage. Body, length about 3 mm, width about i mm ; 65 young
specimens taken from the marsupium of a single female ; shape in dorsal view as in
Fig. 3a. Head with rounded eye lobes and forwardly produced rostrum as seen in
the male ; eyes large. Peraeon tergite i almost twice length of tergites 2-6, anterior
and posterior margin of tergite i broadly rounded, segment 7 reduced and con-
tinuous with pleon, dorsal side of peraeon flattened; no penis lobes evident. Pleon
rectangular, all segments free laterally; telson slightly longer than wide, rounded
posterior margin bearing about 13 long plumose setae.

Appendages ; antenna i (Fig. 3b) rounded in cross section and dilated, 7 segments,
of which 4-7 carry prominent groups of aesthetascs; antenna 2 (Fig. 3b) slender, a
little longer than antenna i, 8 segmented with a small group of terminal setae;
mandible, maxilla i, maxilla 2, and maxilliped similar to those of female except
much smaller and the terminal hooks evident only as small rounded tubercles.
Peraeopods increasing in length very slightly from 1-7, basis and ischium well
developed, merus and carpus short, propodus strong, and dactylus long, pointed,
and sharply curved; peraeopods 1-3, merus with long spine-like seta on antero-
lateral corner (Fig. 3c), also a characteristic number of hooks on inner side of each
peraeopod, peraeopod 1-2 with a pair of hooks on propodus (Fig. 30), peraeopods
3-4 have a pair of large hooks on propodus and a single curved spine on carpus
(Fig. 3d), peraeopod 5 with two propodal and two carpal spines (Fig. 36), peraeopod
6 bears two spines on propodus, three on carpus, and a single one on the merus
(Fig. 31). Pleopods bilobed (Figs 3g-k), leaf -like with long marginal plumose setae,
setae absent from endopods of pleopods 3-5 ; peduncle of pleopods 1-5 each with a

334

R. J. LINCOLN

FIG. 3. Lironeca africana sp. nov. a-1, second marsupial stage: a, body entire, dorsal;
b, antennae i & 2 ; c, peraeopod i, left; d, peraeopod, 3, left; e, peraeopod 5, left; f, perae-
opod 6, left; g, pleopod i, left; h, pleopod 2, left; i, pleopod 3, left; j, pleopod 4, left;
k, pleopod 5, left; 1, uropod & telson entire, m-q, first marsupial stage; m, body entire,
dorsal; n, peraeopod i, left; o, peraeopod 6, left; p, pleopod i, left; q, pleopod 2, left;
bar scale 0-5 mm.

LIRONECA (ISOPODA; CYMOTHOIDAE) 335

set of coupling spines ; exact number of marginal setae on the pleopods is somewhat
variable; uropods (Fig. 3!) with long plumose setae, the exopod bearing a single
large curved spine at the distal tip.

REMARKS. As might be expected in a group which shows marked polymorphism,
there are some small morphological variations in the paratype material. It is
important to appreciate this variability when attempting to identify material such
as this, and the lack of such appreciation has undoubtedly been partly responsible
for the very large number of species which have been described in the past.

The shape of the telson in females shows some variation from that of the holotype,
and may be broad, almost rectangular, with a stright posterior margin. The
peraeon in one of the specimens is asymmetrical, and there are some differences in
proportional lengths of the peraeon tergites. The first tergite of the peraeon is not
always markedly longer than tergites 2-6, but in all specimens the last tergite is
reduced in length. In the holotype the peraeopods are all approximately the same
length, while in the largest female paratype there is a small increase in the length of
the peraeopods from 1-7, reflected in the size of the ischial segment, the length of
the other segments remaining unchanged. In the males it is important to note that
the body may show signs of asymmetry, and there is a wide range of body size, some
specimens being very small.

The parasites were all taken from the mouth cavities of the cichlid fishes Lampro-
logus elongatus Boulenger and Lamprologus pleurostigma Boulenger. The fishes
examined were part of an extensive collection taken from Lake Tanganyika and
deposited in the collections of the British Museum (Natural History). The female
parasite was attached to the tongue of the fish, with the long dactylae of the peraeo-
pods embedded deeply into the tissues of the host. The isopod was situated with its
head innermost, the posterior end of the body being about level with the angle of the
host's jaw. Despite the large size of the parasite the fish was still able to close its
mouth fully. The male parasite was found in close association with the female,
attached either to the side of the mouth or the inside of the gill chamber. Only one
male was found in the presence of each female.

Lironeca africana is readily distinguished from other freshwater African species
by the general outline of the body and the contour of the posterior margin of peraeon
tergite 7. Lironeca expansus is broadly elliptical in outline and the posterior margin
of tergite 7 has a deeply concave contour, with the pleon strongly immersed into the
peraeon. The head is also deeply set into the peraeon and projects very little, if
any, beyond the general outline of the body. Lironeca tanganyikae has an oval out-
line, somewhat similar to africana, but the margin of tergite 7 is broadly convex with
the median part straight. The pleon is only moderately immersed into the peraeon.
In Lironeca africana the body is more elongate, the convex posterior margin of tergite
7 is an even curve, and the pleon is but weakly immersed. The characters men-
tioned above refer to the mature female stage.

The isopods belonging to the family Cymothoidae are protandrous hermaphrodites
and it has been usual practice to base the description of a new species on the adult
female stage. In the case of Lironeca enigmatica, described by Fryer from Lake

336 R. J. LINCOLN

Tanganyika, the fully mature female was not available and the type series consists
of a number of young male specimens varying both in size and in degree of develop-
ment. The presence of setae on the pleopods and uropods indicates that these
isopods were capable of free swimming, and it still remains to be shown whether or
not the adult parasite is to be associated with the same species of fishes from which
the enigmatica material was taken. Bearing this in mind, and appreciating the
marked transformation which takes place when the parasite takes up permanent
residence with the host, it could prove particularly difficult to establish the true
identity of the adult female. To separate enigmatica and africana it is pointless to
look at the male or female of the latter as they are no longer free swimming, and
have lost all traces of pleopodal and uropodal setae. It only remains to compare
enigmatica with the late marsupial stage of africana, which does have a full setal
compliment in preparation for its early free swimming existence. These can easily
be distinguished on the basis of the number of spines and hooks on the peraeopods
and also the shape of the telson.

A close investigation of the characters possessed by the late marsupial juvenile
stage of cymothoids may eventually provide an answer to some of the many problems
which face the taxonomist working with this group. At the present time there are
many genera which are separated on quite unreliable features and, as Fryer points
out, it is quite possible to refer a given specimen to any of a number of genera. This
has led to some genera receiving a large number of species, Lironeca being a good
example, while others have remained monotypic.

The discovery of this parasite in association with cichlid fishes of the genus
Lamprologus revives the question of the origin of the Lake Tanganyika isopod fauna.
The cymothoids found in the Lake have very obvious affinities with marine members
of the group, with which they no doubt share a common marine ancestral group.
The problem is to establish when the isopods made their entry into the Lake and
acquired freshwater status. Three main possibilities exist, although the first of
these to be outlined can be said to have little support from present knowledge of the
geology and biology of the Lakes region of eastern Africa. In a book 'The Tan-
ganyika Problem' published in 1903, Moore discussed the apparent affinities of the
Lake fauna, with special reference to the molluscan groups, and arrived at the con-
clusion that the fauna had evolved from an ancient Jurassic marine stock which
became isolated when the Lake was cut off from the sea. If the fauna originated in
in this way, the transition of the animals to a freshwater existence would have
occurred within the Lake, and the parasites could have made the change in associa-
tion with their fish hosts. But, as has been noted above this theory of the origin
of the Lake fauna and of the Lake itself has now been largely abandoned.

From the evidence of the present distribution of the freshwater parasites it seems
most likely that the isopods entered the Lake from a connection with the Congo
river system. Lake Tanganyika is the only one of the Lakes in eastern Africa which
is known to have the parasites, and the Congo River system is the only one of the
rivers known to have a freshwater cymothoid fauna. If the isopods entered the

LIRONECA (ISOPODA; CYMOTHOIDAE) 337

Lake region at a time before the isolation of Lake Tanganyika itself, one would
expect to find them in some of the other bodies of freshwater, but this is not the case.
It is now generally accepted that Tanganyika was formed sometime during the
Pleistocene, but the question remains whether the isopods reached the Lake soon
after its isolation, or whether they moved in from the Congo through some river
connection in comparatively recent times. The evidence, limited as it is, tends to
support the former possibility.

The two species of parasite found in the Lake for which a final host is known,
tanganyikae and africana, are both associated with cichlid fishes, and show strict
host specificity. This in itself suggests a long association, and the cichlid fishes
represent a rich endemic element in the Tanganyika fauna. The third cymothoid
known from the Lake, enigmatica, was taken from the clupeids Stolonothrissa and
Limnothrissa, but these isopods were still juvenile and capable of free swimming,
and the identity of their final host has not yet been established. Fryer (1968) points
out that these clupeids have marine ancestors and the parasite may have made the
transition to freshwater with the fish, and further suggests that this could represent
a separate invasion of the Lake by the isopods. This cannot be disproved, but it
is worth noting that the fishes in question are pelagic species which aggregate in
large shoals, and are sufficiently abundant to be commercially exploited (Coulter
1970). Fish which move freely about the Lake and undergo marked diurnal migra-
tion would provide ideal dispersal for the juvenile free living stages of a parasite,
and it seems quite probable that they are acting only as temporary intermediate
hosts for the isopods. The fish are preyed upon by numerous carnivorous fish
species, including some cichlids, and this may be the step by which the parasite
reaches its final host. The speculation will remain until the adult enigmatica has
been found, but if the view outlined above is found to be true, and the adult parasite
is associated with a cichlid fish, it will be less of an enigma than the name was
intended to suggest.

The cymothoid, expansus, found widely in the waters of the Congo, is without
doubt a very close ally of the Tanganyika species. It too would seem to be host
specific, on Eugnathichthys eetweldii, as I can find no record of it having been taken
from any other fish, not even from a closely related species of the same citharinid
fish genus which is also common in the Congo river.

The obvious affinities of the freshwater cymothoids in Africa one with another,
and their very restricted distribution, indicates an entry into Lake Tanganyika from
the Congo river system. The strict host specificity of the parasites and the endemi-
city of the host fishes points to an early invasion of the Lake rather than an entry in
more recent times. However, the nature of this brief summary must remain
speculative until a great deal more is known of the isopods of the African continent.

338 R. J. LINCOLN

REFERENCES
BRIAN, A. & DARTEVELLE, E. 1949. Contribution a l'6tude des Isopodes marins et fluviatiles du

Congo. Annls Mus. r. Congo Beige. C, Zoologie 1 (2) : 77-208.

COULTER, G. W. 1970. Population changes within a group of fish species in Lake Tangan-
yika following their exploitation. /. Fish Biol. 2 : 329-353.
DARTEVELLE, E. 1939. Ichtyoxenus expansus, Isopode parasite dulcaquicole. Rev. Zool.

Bot. afr. 33 (i) : 16-17.
FRYER, G. 1965. A new isopod of the genus Lironeca, parasitic on the cichlid fish of Lake

Tanganyika. Rev. Zool. Bot. afr. 71 (3-4) : 376-384.
- 1968. A new parasitic isopod of the family Cymothoidae from clupeid fishes of Lake

Tanganyika a further Lake Tanganyika enigma. /. Zool., Lond. 156 : 35-43.
GROSSE, J. P. 1963. Le milieu aquatique et Ideologic des poissons dans la region de Yan-

gambi. Annls Mus. r. Afr. cent., Sci. Zool. 116 : 113-271.
MIERS, E. J. 1880. On a collection of Crustacea from the Malaysian region. IV. Penaeidae,

Stomatopoda, Isopoda, Suctoria and Xiphosura. Ann. Mag. Nat. Hist. 5 : 457-472.
MONOD, T. 1931. Sur quelques Crustaces aquatiques d'Afrique (Cameroun et Congo). Rev.

Zool. Bot. afr. 21 (i) : 1-36.

MOORE, J. E. S. 1903. The Tanganyika Problem. London.
NAME, W. G. VAN. 1920. Isopods collected by the American Museum Congo Expedition.

Bull. Am. Mus. not. Hist. 43 : 41-108.

Dr. R. J. LINCOLN

Department of Zoology

BRITISH MUSEUM (NATURAL HISTORY)

CROMWELL ROAD

LONDON, S.W-7

EUNICE MANIHINE SP. NOV. (POLYCHAETA:

EUNICIDAE), A MEMBER OF THE FLAVUS-

BIDENTATE GROUP FROM THE WESTERN

EQUATORIAL INDIAN OCEAN

By M. R. LONGBOTTOMt

SYNOPSIS

A new species of the genus Eunice Cuvier, 1817, is described from 421 m depth in the western
equatorial Indian Ocean. The characteristic features are: yellow, bidentate acicular setae, up
to 4 per parapodium; up to 6 acicula per parapodium; branchiae, with few filaments, present
from setiger 8 to 43; smooth occipital tentacles, the median reaching to setiger 24.

INTRODUCTION

THE few surveys of polychaetes carried out in the western equatorial Indian Ocean
have been restricted to intertidal and shallow-water habitats, rarely exceeding 50 m
depth (Crossland, 1904), and little is known of the fauna of the rough bottoms at
greater depths. During the Royal Society Indian Ocean Deep Slope Fishing
Expedition (January/February, 1969), vertical bottom lines were fished from the
FRV Manihine around the islands and banks in this region of the Indian Ocean
(Forster et al., 1970). One of these lines foul-hooked a piece of coral debris at 421 m,
in which was a eunicid worm ; further examination showed this to be an undescribed
species of Eunice.

The genus Eunice Cuvier, 1817, consists of a large number of valid species (approxi-
mately 140) for which the common specific characters and their variations are
discussed in a recent review (Fauchald, 1970). Briefly, these are the form and
colour of the acicular setae (subacicular hooks), the distribution and form of the
branchiae, and the length and form of the occipital tentacles. Combinations of
these and, frequently, other characters serve to distinguish the species, only rarely is
one of these major characters alone sufficient. Major divisions, based on the form
and colour of the acicular setae, were suggested by Hartman (1944), resulting in
four groups of those species for which the relevant details were known. This
system has been expanded to include a greater number of valid species, and, based
on the distribution of the branchiae, five subdivisions have been proposed for each
of seven groups (Fauchald, 1970).

DESCRIPTION

The single specimen of Eunice manihine sp. nov. consists of 127 setigers and is
approximately 75 mm long (9 mm from the tip of the palps to the posterior edge of
the tenth setiger on the dorsal side, following Fauchald, 1970) and 5 mm wide. The

t Dr. Longbottom died August 6, 1971
Bull. BY. Mus. nat. Hist. (Zool.) 21, 8

M. R. LONGBOTTOM

specimen is cylindrical in the anterior region, and becomes flattened towards the
posterior.

The anterior margin of the prostomium is distinctly notched between the palps
(Fig. i). The length of the prostomium is slightly greater than half the breadth.
The prostomium bears a pair of eyes posterior to the gap between the bases of the

5mm

FIGS 1-7. Eunice manihine. i. Anterior end, dorsal view (left outer occipital tentacle
missing). 2. Setigers 14-17 right side, dorsal view. 3. Posterior end, left lateral
and slightly ventral view. 4. Maxillary plates, dorsal view. 5. Setigers 7-10, dorsal
view. 6. Setigers 7-10, ventral view. 7. Setigers 7-10, diagrammatic representa-
tion, dorsal side firm lines.

EUNICE MANIHINE 341

inner and outer occipital tentacles (Fig. i). Five smooth and very long, slender
tentacles are present ; the unpaired median tentacle reaching back to setiger 24, the
inner lateral tentacles reaching setiger 12 and the outer laterals setiger 3. The
first peristomial segment is two thirds the length of the prostomium; the second
peristomial segment is half the length of the first and the same length as the first
setiger. The peristomial cirri are smooth and reach just beyond the posterior border
of the prostomium.

The maxillae are well developed (Fig. 4); maxilla I is falcate; maxilla II has 12
teeth left and n teeth right; maxilla III has 12 teeth left; left maxilla IV has n
teeth and the combined right maxillae III +IV have 13 teeth ; each maxilla V has
i tooth.

1mm

16 17

FIGS 8-17. Eunice manihine. 8-n. Parapodia from right side, anterior view. 8.
Setiger 4. 9. Setiger 34. 10. Setiger 60. n. Setiger 90. 12. Capillary seta from
setiger 90. 13. Comb seta from setiger 90. 14. Falcigerous seta from setiger 34.
15. Acicular seta (subacicular hook) from setiger 34. 16. Aciculum from setiger 34.
17. Aciculum from setiger 60.

342 M. R. LONGBOTTOM

Branchiae are present only in the anterior third of the specimen: from setiger 8
to setiger 43. The branchiae consist of a single filament anteriorly (Fig. i) with a
maximum of 3 filaments between setigers 17 and 22 (Fig. 2), decreasing posteriorly
to a single filament. The filaments are thinner than the dorsal cirri in the branchial
region (Fig. 2). The dorsal cirri are smooth, anteriorly with a stout base and
elongated tip (Fig. 8), posteriorly digitiform (Figs 3 & n). The ventral cirri are
subulate (Figs 3, 6, 8-n). Two pairs of anal cirri are present (Fig. 3), the dorsal
pair long and the ventral pair very short.

In the majority of the parapodia there are, superiorly, between 10-18 long, slightly
limbate, capillary setae with weak striations (Fig. 12), and, towards the posterior,
5-9 comb setae with 14-16 teeth, usually with the outer tooth on one side appreciably
longer (Fig. 13). Inferiorly, there are 7-9 falcigerous setae, the blades of which are
strongly bidentate, with a rounded hood projecting just beyond the tip (Fig. 14).
The head of the shaft is slightly enlarged. The acicula are yellow with slightly
curved tips (Figs 16 & 17). An average of 3 or 4 are present in each parapodium,
with up to 6 in the middle setigers, decreasing to 2 posteriorly. The inferiorly
situated acicular setae (subacicular hooks) are first present, singly, from setiger
27/28, with 3-4 per parapodium from setiger 31 throughout the remainder of the
setigers. The acicular setae are yellow, bidentate, and hooded (Fig. 15).

DISTRIBUTION

E. manihine is known from one specimen found in a piece of coral debris foul-
hooked on a bottom fishing line from 421 m depth, south of Menai Is, Cosmoledo
(10 S, 47 E), in the western equatorial Indian Ocean.

The type specimen (Ref. no. ZB 1971-1) which is preserved in alcohol has been
deposited in the British Museum (Natural History).

DISCUSSION

No single character is sufficient by itself to distinguish E. manihine from other
members of the genus; the separation is based on a combination of the following
characters: yellow, bidentate, acicular setae; branchiae of few filaments restricted
to the anterior third of the body; very long, smooth, occipital tentacles; the form of
the maxillary plates; and the large number of acicula per parapodium. The first
two characters are common to many species of Eunice and are discussed in greater
detail beiow. References in the literature to occipital tentacles reaching to near or
beyond setiger 20 are uncommon : E. antillensis has been recorded with the median
tentacle to setiger 20 (Ehlers, 1887), E. auriculata to setiger 18 (Treadwell, 1901),
E. palauensis to setiger 18 (Okuda, 1937), and E. tibiana (Pourtales, 1863) to setiger
22 (Izuka, 1912). Of these species only E. antillensis is a member of the flavus-
bidentate group. However, the degree of variation within a species is not known,
and to some extent is dependent upon the state of preservation. This also applies
to the form of the tentacles, whether they are wrinkled or clearly articulated. The
presence of as many as 6 acicula per parapodium is very unusual, contrasting with
the more normal number of 1-3 for species of Eunice. The number of teeth on the
maxillary plate is also greater than is usually found in the genus.

EUNICE MANIHINE 343

Adoption of the system of subdivision proposed by Hartman (1944) and Fauchald
(1970) enables comparisons to be made more easily within this large genus. E.
manihine, in possessing yellow, bidentate acicular setae, clearly belongs to the
flavus-bidentate (A) group of Hartman (1944), and from the distribution of the
branchiae (commencing before setiger 10 and not present after setiger 100), to
subdivision i of Fauchald (1970). As the numbers of setigers may vary in worms
of different sizes, it is suggested that the definition of this subdivision should be
expanded to place the emphasis on the branchiae being present in the anterior third
of the body rather than within a definite number of setigers irrespective of the size
of the worm.

Fourteen species are listed by Fauchald (1970) within this group Ai to which
should be added E. semisegregata Fauchald, 1970, and from which should be
removed E. benedicti (Verrill, 1885) a synonym of E. pennata (Miiller, 1776) (see
Pettibone, 1963) and E. norvegica (Linnaeus, 1767). The latter, in having black
bidentate acicular setae, and branchiae present from setiger 3 to within a few setigers
of the anus (Pettibone, 1963), should be placed in the fuscus-bidentate (B) group,
subdivision 2. All the previously described species in the Ai grouping have the
branchiae commencing on setiger 3. Fauchald (1970) has suggested that this is a
constant feature within a species, though small variations have been recorded, as for
example in E. pennata (Pettibone, 1963). Thus, E. manihine is the sole member of
the flavus-bidentate group in which the branchiae commence before setiger 10, but
after setiger 6. Apart from E. manihine, only two species E. megabranchia
Fauchald, 1970 and E. validobranchiata Monro, 1937 in the Ai group have smooth
occipital tentacles, the remainder have articulated or moniliform tentacles. E.
megabranchia (from 894 m depth in the Gulf of California) and E. validobranchiata
(from 1046 m depth off the South Arabian coast) differ only in the length and shape
of the guards of the falcigerous setae and the form of the ventral cirri; the other
characters are very similar and for comparison with E. manihine may be considered
identical. Specimens of these two species of a similar size to the E. manihine
specimen differ in having branchiae with large numbers of filaments (up to 45)
commencing on setiger 3, appreciably shorter occipital tentacles, fewer acicular
setae per parapodium and fewer teeth on the maxillary plates.

Of the eunicid polychaetes reported from East Africa (Crossland, 1904), only one
species of the flavus-bidentate group was reported E. tubifex Crossland, 1904.
This belongs to Fauchald's subdivision 4, in which branchiae are present from after
setiger 10 to the posterior.

The type specimen of E. manihine shows an abnormality in the segmentation of
setigers 8 and 9 (Figs 5 & 6), resulting in a short spiral (Fig. 7) similar to the form
Buchanan (1893) found in specimens of Lumbriconereis impatiens (Lumbrinereis
tetraura}. It was not possible to examine the internal anatomy of the single speci-
men available.

ACKNOWLEDGEMENTS

I am most grateful to Dr. O. Hartman and Professor J. H. Day for confirming the
separate identity of the species, and to Dr. J. D. George and Mr. P. M. David for

344 M - R - LONGBOTTOM

reading the manuscript. Study leave was granted by the University of Newcastle
upon Tyne for the period of the expedition, which was financed by the Royal Society.

REFERENCES

BUCHANAN, F. 1893. Peculiarities in the segmentation of certain polychaetes. Q. Jl

microsc. Sci. 34 : 529-544.
CROSSLAND, C. 1904. The marine fauna of Zanzibar and British East Africa, from collections

made by Cyril Grassland in the years 1901 and 1902. The Polychaeta. Part III. With

which is incorporated the account of Stanley Gardiner's collection made in the Maldive

Archipelago in the year 1899. Proc. zool. Soc. Land. 1 : 287-330.
EHLERS, E. 1887. Report on "the annelids of the dredging expedition of the U.S. Coast Survey

steamer Blake. Mem. Mus. comp. Zool. Harv. 15 : 1-335.
FAUCHALD, K. 1970. Polychaetous annelids of the families Eunicidae, Lumbrineridae,

Iphitimidae, Arabellidae, Lysaretidae and Dorvilleidae from Western Mexico. Allan

Hancock Monographs in Marine Biology 5 : 1-335.
FORSTER, G. R., BADCOCK, J. R., LONGBOTTOM, M. R., MERRETT, N. R. & THOMSON, K. S.

1970. Results of the Royal Society Indian Ocean Deep Slope Fishing Expedition, 1969

Proc. R. Soc. B. 175 : 367-404.
HARTMAN, O. 1944. Polychaetous annelids. Part 5. Eunicea. Allan Hancock Pacific

Exped. 10 : 1-238.
IZUKA, A. 1912. The errantiate Polychaeta of Japan. /. Coll. Sci. imp. Univ. Tokyo 30 (2) :

1-262.
OKUDA, S. 1937. Polychaetous annelids from the Palau Islands and adjacent waters, the

South Sea Islands. Bull, biogeogr. Soc. Japan 7 : 257-316.
PETTIBONE, M. H. 1963. Marine polychaete worms of the New England region. I. Families

Aphroditidae through Trochochaetidae. Bull. U.S. natn. Mus. 227 : 1-356.
TREADWELL, A. L. 1901. The polychaetous annelids of Porto Rico. Bull. U.S. Fish Commn

20 : 181-210.

Dr. M. R. LONGBOTTOM

NATIONAL INSTITUTE OF OCEANOGRAPHY

WORMLEY

GODALMING, SURREY

ORIGINAL DESCRIPTION OF THE GANGETIC

DOLPHNI, PLATANISTA GANGETIC A,
ATTRIBUTED TO WILLIAM ROXBURGH

By G. PILLERI

IN cetological literature the original description of the Gangetic dolphin is usually
attributed to Heinrich Julius Lebeck. In the third volume of "Neue Schriften der
Gesellschaft Naturforschender Freunde zu Berlin" that appeared in 1801, this
author published a short paper entitled "Delphinus gangeticus beschrieben von
Herrn Heinrich Julius Lebeck zu Trankenbar". "Trankenbar" obviously corres-
ponds to the locality "Tranquebar" in southeast Madras.

In the same year, 1801, Volume 7 of "Asiatick Researches" was published in
Calcutta and contained a paper of William Roxburgh entitled "An Account of a
new Species of Delphinus, an Inhabitant of the Ganges".

It is, therefore, virtually certain that the two descriptions of the Gangetic dolphin
were published in the same year, but the reviews contain no definite indication as
to which of the two appeared first.

Roxburgh's paper was republished in 1803 by Maiden and Wilson of London in a
second edition of "Asiatick Researches, printed verbatim from the Calcutta edition".

In recent reports (e.g. Hershkovitz 1966), data taken from literature onPlatanista
gangetica are frequently referred back to the paper published by Roxburgh in 1803,
the date of publication of the London edition.

When G. Cuvier (1823) changed the designation of the genus Delphinus to Platan-
ista, he claimed priority for Lebeck. E. Home (1818), on the other hand, in his
paper on the morphology of the teeth of the Gangetic dolphin, referred to Roxburgh
as the original author. M. F. Cuvier (1836) who, in nearly all other respects, based
his work on the account of his brother (G. Cuvier 1823), expressed a doubt as to
whom to attribute the original work. Referring to the paper of Home, he added a
footnote at the bottom of page 252 that reads as follows: "T.VII, p. 170, pi. Ill
(e"dit. de Londres, 1803, in-4). M. Home dans sa description des dents de ce
dauphin, cite le memoire de Roxburgh comme ayant paru en 1721 dans les me"moires
de la societe de Calcutta. Ces memoires paraissaient-ils alors? L'e"dition de
Londres est une copie de celle de Calcutta". "1721" is obviously a printing error.

When John Anderson (1833-1900) first took up his activities in India he was
staying with his brother, a doctor and botanist, in the house built in the Botanical
Gardens by Roxburgh. On page 418 of the monograph he published in 1878 on
Platanista and Orcaella, Anderson mentioned that Roxburgh claimed to have
published the first description of the Gangetic dolphin, but made no further comment
on the fact.

A comparison of the two papers of Lebeck and Roxburgh reveals a striking

Bull. Br. Mus. nat. Hist. (Zool.) 21, 8

346 G. PILLERI

similitude in the text, both as regards their presentation and the zoological data they
contain. Here, I should like to give a brief summary of the lives of the two authors.

William Roxburgh. Born at Underwood, Craigie, Ayrshire, on June 3rd, 1751,
William Roxburgh became a distinguished botanist. He studied in Edinburgh
under John Cope and it was Cope who procured him an engagement as assistant
surgeon on a ship of the East Indian Company. After several voyages to the
tropics, he was granted a degree of Doctor of Medicine in Edinburgh and from 1776-
1778 worked in the General Hospital in Madras. In 1780, he became full surgeon
and a year later took up a post at Cocanada, in the delta of the Godavery River,
where he was mainly occupied with applied botany. It was during this period that
he undertook his very extensive collection of plants and illustrations of the Indian
flora. In 1793 he was appointed Superintendent of the Botanical Gardens of
Calcutta but four years later he was forced to return to Scotland on account of ill
health. He went back to Calcutta in 1799, but had to leave India again in 1805.
His state of health continued to deteriorate and he sought recovery at the Cape and
in St. Helena, but in vain. He finally returned to Scotland and died in Edinburgh
on February i5th, 1815.

The numerous and very comprehensive studies undertaken by William Roxburgh,
some of which were published after his death by Wallich, constitute a valuable
contribution to the knowledge of the Indian, and more particularly, of the Bengalese
flora (see Lee 1897). He also compiled reports on water insects, silk worms and land
winds. The work on the Gangetic dolphin (1801) falls in a period of Roxburgh's
life already overshadowed by the outbreak of his serious illness.

Heinrich Julius Lebeck. In spite of extensive investigations in a large number of
libraries at home and abroad, I have not been able to uncover any biographical
details on the life of Lebeck. The only reference I found to his life was contained in
the book of G. Cuvier (1823) wno mentioned that he was a "Danish missionary in
Tranquebar". However, this piece of information is obviously inaccurate as Gosch
(1875), in his book on early Danish zoological literature, describes Lebeck as a
German missionary. Furthermore, the biographical lexicon in the royal library in
Copenhagen contains no mention of Lebeck.

The descriptions of Lebeck and Roxburgh

It should be pointed out that Roxburgh stresses the discovery of a new species in
the title of his work, "An Account of a New Species of Delphinus", and Lebeck does
not. Both descriptions refer to a male specimen from the Calcutta region of exactly
the same size (!). What is even more significant is that not only were the measure-
ments quoted in the two papers identical, but in Lebeck's paper British linear
measurements and British avoirdupois weights were used. If the original contribu-
tion had been German one would logically expect the author to have employed
German units. The two reports start with Linnaeus and state that the Gangetic
dolphin is a fifth, new species to be added to the four species of dolphins described
by Linnaeus. The head, teeth, tongue, eyes, external auditory passages, fins and
genitals are described in the same order. Both authors found nematodes and plant
seeds in the stomach. Both mention the thickness and strawyellow colouring of the

THE GANGETIC DOLPHIN PLATANISTA GANGETICA 347

blubber as well as the fact that dolphins oils are used for medicinal purposes by the
natives.

Not only is the order in which the organs are described in the diagnosis of the
species identical, but also, whole sentences are repeated word for word.

A comparison of the texts leads to the conclusion that one of the two authors was
acquainted with the text of the other before he published his own work. It is also
significant that Lebeck mentions a "zweiten mir gezeigten Russel eines solchen
Tieres" 1 (page 282). I presume that this rostrum could only have been shown to
him within the natural science circles of Calcutta, the town where research on the
Gangetic dolphin originated with the collections and observations of Roxburgh,
Blyth and, later, John Anderson.

Here it is worth while noting that in the British Museum (Natural History) a cast
of rostrum Nr. 1884.5.3.1, River Ganges, is exhibited with the label: "the original
specimen now in the Museum of the Royal College of Surgeons 2 was described and
figured by Dr. Roxburgh it exhibits in great perfection the characters of the teeth
in aged animals, which are quite different from these of the young". It is tempting
to conclude that this is the specimen previously shown to Lebeck.

It is not certain that the "Neue Schriften der Gesellschaft Naturforschender
Freunde zu Berlin" was also available in Calcutta in 1801. Certainly no copy
of this journal is included in the Indian Museum library to-day. "Asiatick
Researches", on the other hand, must have been easily accessible to Lebeck who
was living in India, or had lived in India, at about that time.

There is no doubt that Roxburgh made very thorough observations of the Hughly
river dolphins, since his house, as Anderson (1878) explained, resembled the bridge
of a ship and commanded an extensive view of the Hughly river. Gangetic dolphins
still swim in this river to-day as they did in the past. Roxburgh was also interested
in hydrobiological problems (water insects) which is further proof of his familiarity
with Indian rivers. He was considered an expert responsible for many botanical
discoveries and descriptions of Indian plants.

Although he was predominantly a botanist Roxburgh did have the preliminary
training as a surgeon so, like Anderson, he would have had the basic knowledge of
anatomy that would have made him competent to deal with a dolphin. It is incon-
ceivable to me that a man of his experience would need to copy a report of Lebeck.
The contrary is much more plausible. Therefore, I am of the opinion that William
Roxburgh, and not H. J. Lebeck, was the first author to describe this interesting
species of cetacean, and suggest that in the future publications on the Gangetic
dolphin, the species should be referred to as Platanista gangetica (Roxburgh 1801).

ACKNOWLEDGEMENTS

The biographical data on William Roxburgh were placed at my disposal by my
friend and colleague, Dr. Francis C. Fraser, British Museum (Nat. Hist.), London.
Grateful acknowledgements is also made of the kind co-operation of the Trustees and

1 "the second rostrum I was shown of such an animal".

2 The specimen was destroyed by enemy action during the second world war.

348 G. PILLERI

the Keeper of Zoology Dr. J. P. Harding of the British Museum (Nat. Hist.), London,
who allowed me admittance to the osteological collections. I also wish to extend my
thanks to Dr. M. Gihr, Dr. C. Kraus, Prof. B. Horning in Berne, and Dr. U. M0hl,
Zoological Museum Copenhagen, for their help in the bibliographical research and to
Mrs. Diane M. von Nordheim, Geneva, for the English translation. The work was
sponsored by the Swiss National Fund for the Promotion of Scientific Researches.

SUMMARY

The attribution of the original description of Platanista gangetica to Lebeck is
contested. It is first established that the descriptions of Lebeck and Roxburgh
were published in the same year, i.e. in 1 80 1, and the reasons for concluding that the
description of Lebeck was copied from that of Roxburgh are set out. It is suggested
that Roxburgh be considered the author of the original description of the Gangetic
dolphin.

REFERENCES

ANDERSON, J. 1878. Anatomical and Zoological Researches: comprising an Account of the

Zoological Results of the two Expeditions to Western Yunnan in 1868 and 1875; and a

Monograph of the two Cetacean Genera Platanista and Orcella. 2 vols. London (B

Quaritch) .
CUVIER, M. F. 1836. De 1'histoire naturelle des Cetaces ou recueil et examen des faits dont se

compose 1'histoire naturelle de ces animaux. 416 pp. Paris (De Rovet).
CUVIER, G. 1823. Recherches sur les Ossements Fossiles de Quadrupedes, ou Ton retablit

les characteres de plusieurs especes d'Animaux que les revolutions du Globe paraissent avoir

detruites &c. 5 : 279-280. Paris (Dufour et D'Acagne).

GOSCH, C. C. A. 1875. Udsigt over Danmarks Zoologiske Literatur. 2. Afd., Bd. 2, Kobenhavn.
HERSHKOVITZ, P. 1966. Catalog of Living Whales. Bull. U.S. natn. Mus. 246 : 1-259.
HOME, E. 1818. A description of the teeth of the Delphinus gangeticus. Phil. Trans. R.

Soc. 108 : 417-418.
LEBECK, H. J. 1801. Delphinus gangeticus beschrieben vom Herrn Heinrich Julius Lebeck zu

Trankenbar. Neue Schr. Ges. naturf. Fr. Berl. 3 : 280-282.

LEE, S. 1897. Dictionary of National Biography. 44 : 368. London (Smith, Elder & Co.).
ROXBURGH, W. 1801. An Account of a new Species of Delphinus, an Inhabitant of the

Ganges. Asiat. Reschs. 7 : 170-174. Calcutta.
1803. (printed verbatim from the Calcutta edition). London.

Professor G. PILLERI
BRAIN ANATOMY INSTITUTE
UNIVERSITY OF BERNE
3072 OSTERMUNDIGEN
SWITZERLAND

A NEW GENUS AND SPECIES OF SUDAN

LEECH FORMERLY CONFUSED WITH
LIMNATIS NILOTIC A (HIRUDINIDAE S.L.:

HIRUDINEA)

By L. R. RICHARDSON

SYNOPSIS

A new genus is based on a leech from Zalingei Swamp, Sudan. It has 3 pairs of narrow
broken lines on the dorsum, somitally repetitive supramarginal maculations, and a marginal
light stripe; but there are 16 complete 5 -annulate somites, the posterior sucker is of moderate
size, ejaculatory bulbs are present, and there are linear somital sense organs on the dorsal aspect
of the posterior somites. A closely similar leech is recorded also from the Nile, near Fashoda.

INTRODUCTION

LEECHES with salivary gland papillae on the jaws have been found in all regions,
essentially between 35 North and South. Of these, only leeches in the Mexican
genus Limnobdella Blanchard 1893, the Australian genus Quantenobdella Richardson
1969, and the present leech have 16 complete 5-annulate somites, all others (Soos,
1969) have 15 such somites, including the leeches in the Ethiopian genus Limnatis
Moquin-Tandon 1826 based on L. nilotica (Savigny 1822). The genus Limnobdella
(v. Richardson, 1969 : 106, 'Potamobdella'} has a macrobdelloid pharynx with some
muscular ridges ending independently between the jaws on the entrance to the
pharynx, multiple small testes in each somite, the epididymis formed on both limbs
of a simple primary loop on the anterior region of the male paired ducts, no ejacula-
tory bulbs, the median regions bimyomeric and the female median region with an
acaecate vagina and vaginal duct. The genus Quantenobdella described as lacking
salivary gland papillae and having an hirudoid pharynx, is now found to have
minute papillae and the pharynx smooth internally. Quantenobdella has simple
saccular testes; the anterior region of the male paired duct lacks a loop and the
epididymis is entirely posterior to the ejaculatory bulb, the two being linear in
relationship; the median regions are bimyomeric and the female median region has
a caecate vagina and a vaginal duct.

In the present leech, the pharynx is hirudoid; the testes are simple, saccular; the
anterior region of the male paired duct forms a simple primary loop with the epidi-
dymis on the initial limb and an ejaculatory bulb on the terminal limb in a sub-
parallel relationship; the median regions of the reproductive systems are as in
Limnobdella.

These differences warrant the provision of a separate and new genus for the leeches
from the Sudan. It has been shown (Richardson, 1969) that the content of the
former family Hirudinidae based on the genus Hirudo was unacceptably hetero-
genous. Although separate families were provided then for Australian, Nearctic

Bull. Br. Mus. nat. Hist. (Zool.) 21, 8

350 L. R. RICHARDSON

and Neotropical genera, the provision of a family for Oriental and Ethiopian leeches
having an acaecate vagina, as in the present leech, must be postponed until the aquatic
jawed sanguivores of these regions are more adequately known. For the time being,
the new genus can be referred to the Hirudinidae s.l.

Moore (1939) identified leeches in single collections from the American Museum of
Natural History, the United States National Museum, and the British Museum
(Natural History), all as being Limnatis nilotica. He briefly referred to them as
resembling this species in having the annulation in no way differing from published
accounts and with small papillae, large caudal suckers, the morphology of the crop
caeca and of the reproductive systems as in Moquin-Tandon (1846, pi. vi, Haemopis
sanguisuga) and Dequal (1912). The characteristics as taken from these authors are :
a small anterior pair and a larger posterior pair of caeca on each compartment of the
crop ; the epididymis formed on both limbs of a simple primary loop on the anterior
region of the male paired duct which lacks ejaculatory bulbs; the two ejaculatory
ducts enter independently into the atrium (Moore, 1939, Fig. 58) and the vagina
acaecate with a vaginal duct of the same length as the vagina. The vaginal duct is
shown as short by Dequal (1912, Fig. 13) and the vagina possibly caecate ; but there
is definitely no duct according to Moquin-Tandon (1846, pi. vi, Figs 15, 17). Moore
recorded from the specimens before him, the presence of only 45 to 60 teeth, a
pattern of 3 pairs of continuous or broken dark dorsal 'lines', yellow marginal stripes
and the venter immaculate as in 'typical North African' examples of this species.

Moore's material in the British Museum (Natural History) was collected in 1925
and came from Zalingei Swamp, former British Sudan. Specimens from this collec-
tion and another from the Nile near Fashoda, 1907, were loaned to Keegan et al.
(1969) who figure without other description a leech from one of these collections,
under the name of Limnatis nilotica. (The external features, Figs 7A&B, 8A&B;
the jaw, Figs 8 C & D, and reproductive system, Fig. 8 E.)

The reproductive system as shown in Keegan et al. differs from the illustration in
Moore (1939, Fig. 58) of the system in a specimen in the U.S. National Museum
collection No. 5501, from between Abyssinia and British East Africa, in that Keegan
et al. show small ejaculatory bulbs embracing the epididymi and the two ejaculatory
ducts joining to form a distinct common duct leading to the atrium. No such
common duct has been known in jawed sanguivorous or macrophagous leeches, the
two ducts always entering the atrium independently.

Re-examination of specimens from the 1925 Zalingei Swamp and 1907 Fashoda
collections confirm the basic pattern as described by Moore and shown in Keegan
et al. The jaws carry minute teeth and papillae. Three dissected leeches show
anterior and posterior paired caeca on the crop compartments but differ from Keegan
et al. in having the ejaculatory ducts entering independently into the atrium, as in
Moore (1939). They further agree with Moore in having a single primary loop on
the anterior region of the male paired duct but differ by having the epididymis on
the initial limb of the loop subparallel to the bulb on the terminal limb, as in Keegan
et al. They differ again from Moore in having linear somital sense organs on the
dorsum of the posterior part of the body, as in most species of Limnatis (i.e. excepting
paluda and possibly nilotica).

A NEW GENUS AND SPECIES OF SUDAN LEECH 351

In having 16 complete 5-annulate somites, the specimens differ from the genus
Limnatis as defined by Moore (1927) and Soos (1969). As I have shown previously,
if we attempt to follow Moore (1927 et seq.) and admit such a difference into a single
genus, the generic entity fails on other grounds (Richardson, 1969, 1970). There is
no established genus suitable for them. Accordingly I provide a new genus as
below.

DESCRIPTION

Aliolimnatis gen. nov.

Hirudinidae s.l. with a monostichodont condition ; 16 complete 5-annulate somites
(ix to xxiv) ; xxv, 4-annulate ; somital sense organs, large and obvious on the dorsum,
circular anteriorly, mostly linear and oblique posteriorly; jaws, moderate in size;
teeth, minute, about 80; salivary gland papillae on the jaws; dorsal salivary glands,
compact, a single mass without obvious columns of aggregated ducts; radial muscles,
a distinct extrinsic system in viii and ix ; pharynx and associated structures, hirudoid ;
mouth and lumen of pharynx, narrow, the lumen tubular; pharynx with six internal
muscular ridges as dorsomedian and ventrolateral pairs, each pair joining to enter a
jaw, none ending independently between the jaws; pharynx terminating posteriorly
in ix; crop compartments in x to xviii each with an anterior small and posterior
larger pairs of caeca, the posterior pair in xix forming the postcaeca extending to
xxvi; intestine, simple tubular, joining terminally to the rectum; genital pores in xi
and xii bs/be; testes, normally 10 pairs; anterior region of male paired duct folded as
a simple primary vertical loop, the epididymis on the initial limb posterior to the
ejaculatory bulb on the terminal limb, the relationship subparallel; median regions,
bimyomeric, mesomorphic; penis sheath reflected anteriorly; oviducts, short, of the
length of the ovaries; atrium, thick-walled, large; common oviduct, thick-walled,
longer than the oviducts, continuous with the acaecate vagina; vaginal duct, short.

Size, (?) medium. Pattern, longitudinal interrupted narrow dark bands, marginal
contrast stripes.

Aquatic. Sanguivorous. Ethiopian Region.

The name Aliolimnatis is derived from alms another, and limnatis of the marshes.
The gender is feminine.

TYPE SPECIES: Aliolimnatis diver sa sp. nov.

Aliolimnatis diversa sp. nov.
Fig. i A-F

HOLOTYPE: B.M.(N.H.) Reg. No. 1970 3 i

SCHIZOHOLOTYPE : B.M.(N.H.) Reg. No. 1970.3.2 right ventrolateral jaw (micro-
slide).

PARATYPES: B.M.(N.H.) 1970.3.3 (18-0 mm long). 1970.3.4.
LOCALITY: Zalingei Swamp, Sudan. (Coll. Admiral Lynes.)

352 L. R. RICHARDSON

GENERAL FORM. Holotype (Fig. i, E.) preserved in alcohol, rather strongly
contracted with the annuli generally at least as high as long; short, heavy-bodied,
depressed, the dorsum almost flat, the margins obtusely rounded, the venter flat,
and the depth nearly uniform along the length of the body.

The anterior sucker broad, the aperture transverse ; the velum proper contracted,
thick margined and turned ventrally; the body widening gradually behind the
sucker, at first subcircular in section, then widening more rapidly to the anterior end
of the clitellum and from here to the post-nephridial region, the width uniform, the
margins parallel, converging abruptly in the post-nephridial region to form the
narrow base to the sucker. The posterior sucker is about half the maximum width
of the body.

Total length, 22-0 mm; depth generally 3-0 mm; width at iv/v, 2-4 mm, at vi/vii
4-0, at ixjx 4-8; clitellum and testicular region, 6-0 mm wide; basis for sucker, 2-0
and diameter of sucker 3-0 mm.

COLOUR AND PATTERN. (Fig. i, A, B, C, E.) Preserved, faded; general back-
ground colour, faintly yellowish brown on the dorsum, paler on the immaculate
venter which is separated by a pair of dark longitudinal bands from distinct pale
cream marginal stripes. A dark patch across the posterior quarter of the dorsum of
the sucker which is otherwise pale above and below.

Three pairs of weakly indicated maculated interrupted dark longitudinal lines
form narrow bands on the dorsum, dividing it into a median stripe, a wide inner and
two narrow outer pairs of longitudinal stripes, all of the background colour so that
the dorsum is not brilliantly or colourfully striped, and the marginal stripes provide
the only contrast colour other than the dark bands. The maculations which form
the dark bands have the appearance of very large individual chromatophores.

The ocular arch is on the edge of the background colour, with the 5th pair of eyes
in from this edge which becomes lateral to the supramarginal line in viii and immedi-
ately lateral to the line of marginal somital sense organs along the greater length of
the body including xxvi, so defining above the cream marginal stripe continuous
around the velum and back to xxvi/xxvii.

The inner and middle pairs of dark bands extend along the medial and lateral
borders of the paramedian fields; the inner pair commencing in viii and extending
to xxvi a.2, define the median light stripe between these limits as filling the median
field and including the paramedian sense organs. Between the ocular arch and viii,
the median field and much of the interocular area are vaguely darker than the
general colour elsewhere. The middle pair of narrow bands extend from in vii into
xxvi (possibly also into xxvii for the dark patch on the sucker is divided as though
into topographic equivalents of the inner and middle pair of bands). Between the
inner and middle paired bands, the inner paired light stripe fills the greater part of
the paramedian field, increasing in width as this field widens along the length of the
body. The outer paired narrow dark bands are lateral in the intermediate field,
extend from in viii back into xxv and between these limits define the middle paired
stripes which include the intermediate sense organs and the greater part of the
intermediate field.

A NEW GENUS AND SPECIES OF SUDAN LEECH 353

The narrow outer paired stripes are lateral to the outer paired bands, denned
laterally by the marginal stripe, and include the supramarginal sense organs, the
supramarginal field and marginal sense organs along the greater length of the body.

In the middle half of the body, there are distinct small darker patches close to the
lateral edge of the outer paired stripe. Each patch is restricted to an annulus.
These occur with recognizable regularity on bi and bs, in some somites on b 2 and b$
and in a few somites on bi on only one side or the other. Such patches are present
from viii to xxvi. The patches are spaced, nowhere give the appearance of a band,
and form a distinct subsidiary pattern as in Hirudinaria and Poecilobdella.

The dark band separating the marginal stripe from the venter, extends along the
line of the submarginal sense organs and briefly into the fields on either side.

ANNULATION. (Fig. i, A, B, C.) Preserved, contracted. Intersomital and
interannular furrows, very deep, equivalent; no obvious division of annuli into
couplets or triplets; somital limits not indicated generally; annuli liberally divided
by fine longitudinal lines into small rectangles. Somital sense organs are circular
and difficult to detect on the venter, but obvious in large clear patches on the dorsum
where all are circular in the anterior region but progressing posteriorly: the inter-
mediates are elongate, linear and oblique to the long axis of the body; the supra-
marginals are linear along the long axis ; then the paramedians are linear and oblique ;
and the supramarginals, linear and transverse. Linear somital sense organs occur
back into xxvi ; all are circular in xxvii. Sensillae are obvious with a central sensilla
surrounded by others forming a rosette, the rosettes arranged as a row across the
annulus, each rosette situated in a rectangle. Nephropores are obvious, minute, and
situated close to the posterior border of ai and b 2 just laterally to the line of the
intermediate sense organs.

The velum proper is broad with a thick incised margin. The velum includes the
ist to 3rd pairs of eyes ; the first furrow iv/v does not reach the margin and the dorso-
lateral lobe of the margin of the sucker is not strongly defined anteriorly; v, 2-
annulate above, aia2 with the first eyes and first detectable paramedians > as, the
furrow aia2/aa reaching into the supramarginal fields with aia2 briefly forming the
lateral margin of the sucker which is completed by uniannulate v which also forms
the ventral margin; vi is 3-annulate above, the 5th pair of eyes in a2, ai<a2>as,
ai/a2 ending in the supramarginal field and vi is 2-annulate below with aia2 > as;
vii, 3-annulate above and below, ai<a2<as; viii, 4-annulate with ai (=vii as) >
a 2 > bs = be and with the first pair of nephropores on ai ; ix to xxiv are all 5-
annulate (total 16) ; ix, bi = b 2 = a 2 < b 5 = b 6 ; x, bi = b 2 < a 2 = b s > b e , as
also xi; due to contraction, the relative lengths of annuli are not assessable with
confidence in xii to xvii; xviii, bi = b2 < a2 < bs > be, as also xx and xxi, so that
b 5 may possibly be the longest annulus in typical 5-annulate somites of the middle
nephric group; xxiv, bi = b 2 < a 2 > b 5 = be, and the last nephropores on b 2 ;
xxv, 4-annulate above and below, bi =b 2 < a 2 < a 3 ; xxvi, 3-annulate above,
ai < a2 > as but definitely 4-annulate below, ai being divided into bi < b 2 by a
well-formed furrow which does not quite reach the margins of the body and is not
to be seen from above; xxvi 3.3 is the last annulus across the venter and is much

354

L. R. RICHARDSON

2.0

FIG. i. Aliolimnatis divevsa gen. et sp. nov. Dorsal annulation and pattern A. somites i
to x, B. midnephric region, and C. somites xxv to ^.wn and posterior sucker. D. Jaws,
and internal muscular ridges of the pharynx opened along the midventral line (arrow
marks mid-length of pharynx) . E. Ventral aspect showing general form and submarginal
bands. F. Anterior region of male paired ducts, male median region, and female repro-
ductive system (arrow marks dorsal aspect of male primary loop) . All figures from the
type. Roman numerals indicate somites and somital ganglia. Somital ganglia
represented at relative size. A. gl. albumin gland; at. atrium; c. od. common oviduct;
ej. b. ejaculatory bulb; epid. epididymis; ov. ovary; p.s. penis sheath; pr. prostate; va.
vagina; va. d. vaginal duct; v.d. vas deferens. Scales in mm, 0-5 mm unless otherwise
indicated.

A NEW GENUS AND SPECIES OF SUDAN LEECH

355

shortened; xxvii, incomplete 2-annulate with a brief accessory annulus. The anus
is at the posterior border of xxvii.

Dorsum of the posterior sucker with 6 concentric furrows; some linear sense
organs are detectable. The ventral face has some 20 primary muscle bands centrally,
dividing to give in the order of 50 at the margin.

ALIMENTARY TRACT. (Fig. i, D.) The jaws are monostichodont, compressed,
the dorsomedially wider than high at the median end with the base 0-6 mm wide,
and the jaw 0-4 mm high; the dental margin is so low convex as to appear almost
straight and is 0-9 mm long. There are about 79 teeth on the right ventrolateral
jaw. The teeth are narrowly spaced, minute, about 0-018 mm tall at the median
end, diminishing very gradually along the row so that teeth in the middle of the row
are still 0-015 mm tall. The dorsomedian jaw is housed in an open groove, the
ventrolaterals in open pits and the margins of all are so poorly denned that the pits
and groove appear non-morphological. The salivary gland papillae are in three
rows, small excepting the basal row which has some larger papillae. The dorsal
salivary glands form a compact mass in vii to ix with no indications of columns of
aggregated ducts. The extrinsic radial musculature of the pharynx is sparse but
forms an obvious system in viii and ix. The pharynx commences at vi/vii. The
entrance to the pharynx is narrow, barely wider than the base of the dorsomedian
jaw, and the lumen is more simply tubular than tapering. The pharynx has a thin
muscular wall with six internal muscular ridges arranged as dorsomedian and ventro-
lateral pairs, each pair fusing before entering the base of the corresponding jaw,
none ending independently on the margin of the entrance to the pharynx. The
pharynx terminates in the posterior portion of ix followed by a short simple com-
partment in ix as the first portion of the crop which has a short small compartment
in x with an anterior small and posterior larger pairs of lateral caeca, as also on xi
to xviii in which the compartments increase progressively in length and breadth, the
anterior caeca remaining simple, small, and the posterior caeca increasing relatively
in size, extending into the paramedian chamber and posteriorly to lie laterally to the
anterior caeca of the following somite. In xix, the anterior caeca are small and
restricted to the median chamber, the posterior pair form elongate postcaeca reflected
in the paramedian chamber, reaching to xxvi and subdivided laterally into lobes. A
short length of the crop completes the compartment in xix, connecting terminally at
xix/xx to the simple tubular intestine which tapers to xxiv/xxv where it enters
terminally into the short simple rectum.

REPRODUCTIVE SYSTEM. (Fig. i, F.) The genital pores are at xi and xii
The testes are simple saccular. There are normally n pairs situated inter-
somitally at xiii/xiv to xxiii/xxiv. The vasa deferentia extend in the paramedian
chambers to the level of xi\xii with the anterior region of each male paired duct
folded vertically in a simple primary loop in xi. The small epididymis is tortuous
on the initial posterior limb of the loop with the muscular ejaculatory bulb on the
anterior terminal limb, and the relationship of the organs is sub-parallel. The
ejaculatory bulbs are subfusiform. Muscular ejaculatory ducts extend medially
into the median chamber to enter independently into the dorsal aspect of the male

356 L. R. RICHARDSON

atrium, not the ventral aspect as is usual, because the male median region is formed
on an anteriorly directed primary loop reflecting at x/xi so that the atrium is at the
posterior end of the procurrent limb continuous with the muscular penis sheath
which has the procurrent limb dorsal to the terminal recurrent limb.

The ovaries are elongate and tapering. The short oviducts join just behind
ganglion xii to form an unusually large thick-walled atrium lined with a longi-
tudinally rugose epithelium. The atrium tapers into the thick- walled common
oviduct which is slightly tortuous and continues into the acaecate vagina. The
vagina is sharply differentiated from the strongly muscular short thick-walled
vaginal duct.

The prostate is a very large thick cap covering the atrium and extending briefly
along the procurrent limb of the penis sheath. The albumin glands are a thick
investment of the atrium and of most of the common oviduct.

MORPHOLOGY OF THE PARATYPES AND OTHER MATERIAL. Four specimens rang-
ing from 16-0 to 25-0 mm in length. The general form is as in the holotype. The
diameter of the posterior sucker is slightly wider than half of the maximum body
width, av. 56 % (range, 50 % to 65 %) and most doubtfully of the width of the body
excepting in extreme extension of the animal. Colour, as the holotype. Pattern,
as the holotype in three specimens with the paired dark bands more pronounced,
and all are distinctly maculate, nowhere continuous. The fourth specimen (25 -o mm)
has only scattered sparse spaced maculations which do not conform to the topo-
graphy of the paired bands in the others. Dorsal somital sense organs are distinctly
linear on the posterior region of the first three specimens, but very vaguely indicated
on the fourth (25-0 mm). Annulation, as in the holotype, excepting in the fourth
specimen (25 -o mm) in which ii\iii and iii/iv cross the paramedian and median fields.
In all, there are 16 complete 5-annulate somites, and xxvi is 3-annulate above and
4-annulate below.

The differences indicate a possible second species represented by the 25-0 mm
specimen.

Two SPECIMENS FROM THE NILE, NEAR FASHODA. Material from the second
collection available to me from the British Museum (Natural History) included two
specimens. Data on the label are: Limnatis nilotica Savigny. 1907.11.12 1/3.
Loc : Nile near Fashoda. Pres : Loat Collection.

The specimens are 39-0 and 49-0 mm long. Preserved, faded, general colour
much as in A. diversa but slightly darker. Essential pattern as in A. diversa,
excepting the paramedian field is moderately filled with many dark brownish bold
maculations which are erratically placed and blend into the medial and lateral paired
bands of this field. This gives the appearance of a broad band across the paramedian
field and accentuates the median stripe so making it a prominent feature of the
dorsum. This is much as shown in Keegan et al. (1969, Fig. 7).

Somital sense organs obscure, but some linear oblique organs are detectable
posteriorly on the dorsum; sensillae, arranged as a rosette.

Annulation, as in A . diversa, with ix to xxiv complete 5-annulate (total 16) except-
ing that xxiv a.% is incomplete briefly in the ventral median field, xxvi is simply

A NEW GENUS AND SPECIES OF SUDAN LEECH 357

3-annulate above and below with the annul! crossing the. venter as thin cutaneous
folds, and xxvii, 2-annulate without an accessory annulus.

Alimentary tract as in A. diver sa, excepting the intestine joins subterminally to
the rectum so that there is a distinct rectal appendix about 2\ times as long as wide.
The jaws were not studied for the dentition.

Reproductive system, 39-0 mm specimen. General morphology and morpho-
logical relationships as in A. diversa, the male median region reflecting at ganglion
x; excepting that the anterior region of the male paired duct forms a posteriorly
directed loop from xi into xii, on which the epididymis is on the recurrent limb and
ventral to the bulb on the procurrent limb, the relationship is subparallel.

Annulation and pattern suggest another possible separate species in Aliolimnatis,
a question to be decided only from the study of more and preferably better material.

ACKNOWLEDGEMENTS

I am greatly indebted to Mr. R. W. Sims, Annelida Section, Department of
Zoology, British Museum (Natural History) for the privilege of having this material
for study; to Miss E. Pope, Australian Museum, for help in many ways; to Professor
Marvin C. Meyer, the University of Maine, for his assistance with difficult literature ;
to the Librarian, University of New England, for help with other literature, to the
Science and Industry Endowment Fund for the loan of microscopic equipment.

The study was collateral to researches on Australian Hirudinea assisted by a
grant from the Nuffield Foundation.

REFERENCES

BLANCH ARD, R. 1891. Courtes notices sur les Hirudin6es. i. Sur la sangsue de cheval du

nord 1'Afrique. Bull. Soc. zool. Fr. 16 : 218-221.
DEQUAL, L. 1912. Contributo alia conoscenza degli Irudinei italiani. Arch. zool. ital. 5 (i) :

1-14.
KEEGAN, Hugh L., SEIICHI TOSHIOKA & HIROSHI SUZAKI. 1969. Blood sucking Asian leeches

of families Hirudidae and Haemadipsidae. U.S. Army Medical Command, Japan. 406^

Med. Lab. Spec. Kept. pp. 1-130.
MOORE, J. P. 1927. Arhynchobdellae. pp. 97-302 (in) Harding and Moore, Hirudinea.

Fauna of British India. London.
- 1939. Additions to our knowledge of African leeches (Hirudinea). Proc. Acad. nat. Sci.

Philad. 90 (1938) : 297-360.
MOQUIN-TANDON, A. 1846. Monographic de la famille des Hirudinees. Ed. 2., pp. 1-448 -f

Atlas. Paris.
RICHARDSON, L. R. 1969. A contribution to the systematics of the hirudinid leeches with

description of new families, genera and species. Acta zool. hung. 15 (1-2) : 97-149.
1970. Bassianobdella victoriae gen. et. sp. nov. (Hirudinoidea : Richardsonianidae) .

Mem. Natn. Mus. Viet. 31 : 41-50.
Soos, A. 1969. Identification key to the leech (Hirudinoidea) genera of the world, with a

catalogue of the species, v. Family Hirudinidae. Acta. zool. hung. 15 (1-2) : 151-201.

Dr. L. R. RICHARDSON
4 BACON STREET
GRAFTON, N.S.W.
AUSTRALIA

THE TYPE-SPECIES OF THE GENERA
PHOXINELLUS, PSEUDOPHOXINUS AND
PARAPHOXINUS (PISCES, CYPRINIDAE)

By ETHELWYNN TREWAVAS

Phoxinellus Heckel, 1843 : 1040

Ax this place Heckel listed two species, both new, P. zeregi and P. alepidotus. P.
alepidotus was described on the same page; it had already been named on p. ion,
as an example of species having a certain type of pharyngeal teeth. P. zeregi was
first described on p. 1063. If we were obliged to rely on this publication it would
be allowable to have recourse to page-priority, regard P. zeregi as a nomen nudum
on p. 1040 and adopt P. alepidotus as type-species.

Bleeker early (i86oa : 423-4) made a formal decision in this sense; he denned
Phoxinellus and referred to P. alepidotus as the typical European species ("de
typische 1 europesche soort") ; he noted that P. zeregi should be placed in a separate
genus, for which he proposed the name Pseudophoxinus.

In the same year Bleeker (i86ob) published a "Conspectus systematis Cypri-
norum" in which, though citing no type-species, he defined these two genera in
terms consonant with his designation of the respective type-species in i86oa.

In 1863 Bleeker deliberately changed his mind and designated Phoxinellus zeregi
as "spec, typ." of Phoxinellus, citing Pseudophoxinus Bleeker as a junior synonym.
This appears in three publications of 1863 (ig63a; 31; i863b: 263; i863c: 209)
and in each he proposed a new genus, Paraphoxinus, with P. alepidotus as type-
species.

This is of course inadmissible and Jordan (1919) in 'The genera of fishes' came to
this conclusion without citing all the evidence. He in fact recorded the proposal of
Pseudophoxinus as in the publication here cited as i86ob, noting that no type-species
were named in this brief conspectus and stating that they were supplied in the
"Atlas" of 1863. He seems to have overlooked our 'i86oa' and i86ob is wrongly
given the date 1859 (which belongs only to the first half of the same volume).
Perhaps Jordan was relying on Bleeker's own confession, "Blkr. ol." (Bleeker,
formerly), in the publications of 1863. I have not been able to discover in the works
of either Heckel or Bleeker any reason for Bleeker's change of mind.

The present note, then, confirms Jordan's view, by giving the evidence, that the
only acceptable conclusion is the following :

Phoxinellus Heckel, 1843, type-species P. alepidotus Heckel designated by

Bleeker, 1860; synonym Paraphoxinus Bleeker 1863.

Pseudophoxinus Bleeker, 1860, type-species by monotypy and original designa-
tion Phoxinellus zeregi Heckel = Pseudophoxinus zeregi.

1 This can only mean 'type-species'. The other sense of 'typical of the European spp.' could only be
assumed if other European species had been known at that time, and this was not so.
Bull. Br. Mus. nat. Hist. (Zool.) 21, 8

360 ETHELWYNN TREWAVAS

Bleeker in his three relevant statements of 1863 did not give the reference to his
earlier (1860) designations and his second thoughts have unfortunately been followed
by most authors, among them Steindachner (1882), Pellegrin (1923), Berg (1932,
Zoogeographica), Karaman (1924), Tortonese (1938), Steinitz (1953) and Ladiges &
Vogt (1965). Only Giinther (1868) used Pseudophoxinus as a subgenus (of Leuciscus)
with the single species 'Leuciscus' zeregi, but he banished Phoxinellus altogether by
using Paraphoxinus for P. alepidotus and putting Phoxinellus in the synonymy of
both Leuciscus and Paraphoxinusl

The taxonomy of these little circum-Mediterranean Cyrpinidae is very unsettled,
and authors who have recognized P. alepidotus and P. zeregi as type-species of
distinct genera have not agreed on the limits or definition of the genera, either with
regard to each other or in relation to Pararhodeus Berg and Rutilus (especially R,
tricolor (Lortet); but see Banarescu, 1960 : 67 and 119). The type-species of
Phoxinellus, P. alepidotus, belongs to a group of populations characterized by various
degrees of reduction of the squamation, associated with life in water whose course
runs now in the open, now underground in limestone country in Yugoslavia. Even
the specific boundaries are in doubt and the generic value of the reduction of scales
is questionable. The problem has been stated by some of the authors mentioned
above.

To avoid misunderstanding I should say that I have no plans to work towards a
solution myself. The need to consider P. zeregi in another context prompted me
to look into the nomenclature.

REFERENCES

P. 1960. Einige Fragen zur Herkunft und Verbreitung der Susswasserfischfauna

der europaisch-mediterranen Unterregion. Arch. Hydrobiol. 57 : 16-134, Suppl. maps and

table.
BERG, L. S. 1932. Obersicht der Verbreitung der Susswasserfische Europas. Zoogeographica

1 : 107-208, pi. ii.
BLEEKER, P. i86oa. Ordo Cyprini Karpers. Verh. natuurk. Ver. Ned. Ind. 7 (deel II

no. 5) : 1-492.

i86ob. Conspectus systematis cyprinorum. Natuurw. Tijdschr. Ned.-Ind. 20 : 421-441.

i863a. Atlas Ichth. 3 : 31.

i863b. Notice sur les noms de quelques genres de la famille des cyprinoides. Ver si.
Meded. K. Akad. wet. Amst. 15 : 261-264.

i863c. Systema cyprinoideorum revisum. Ned. Tijdschr. Dierk. 1 : 187-218.
GUNTHER, A. 1868. Cat. Fish. Brit. Mus. VII.

HECKEL, J. J. 1843. Ichthyologie in Russegger's Reisen vol. i : 993-1250.
JORDAN, D. S. 1919. The genera of fishes. Part 2 (pp. i-ix, 163-284, i-xiii).
KARAMAN, S. 1924. Pisces Macedoniae. 90 pp. (Split).
LADIGES, W. & VOGT, D. 1965. Die Susswasserfische Europas. 250 pp. (Hamburg and

Berlin. Paul Parey).
PELLEGRIN, J. 1923. Poissons de la Syrie rapportes par M. Henri Gadeau de Kerville in

Voyage zoologique d' Henri Gadeau de Kerville en Syrie (Avril-Juin 1908) : 5-37, pis. i-v.
STEINDACHNER, F. 1882. Beitrag zur Kenntniss der Fische Afrikas und Beschreibung einer

neue Paraphoxinus-Kri. aus der Herzegowina. Denkschr. Akad. Wiss. Wien 45 : 1-18.

PHOXINELLUS, PSEUDOPHOX I N US AND PARAPHOXINUS 361

STEINITZ, H. 1953. The freshwater fishes of Palestine. Bull. Res. Counc. Israel 3 : 207-227.
TORTONESE, E. 1938. Viaggio del Dott. Enrico Festa in Palestina e Siria (1893). Pesci.
Boll. 1st. Mus. Zool. Univ. Torino 46 (3) no. 85 : 1-48, pis. i and ii.

Miss ETHELWYNN TREWAVAS D.Sc.

c/o Zoology Department

BRITISH MUSEUM (NATURAL HISTORY)

CROMWELL ROAD

LONDON, S.W-7

A LIST OF SUPPLEMENTS
TO THE ZOOLOGICAL SERIES

OF THE BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

1. KAY, E. ALISON. Marine Molluscs in the Cuming Collection British Museum
(Natural History) described by William Harper Pease. Pp. 96; 14 Plates.
1965. (Out of Print.) 3.75.

2. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier and
Valenciennes. Pp. 180; n Plates, 15 Text-figures. 1967. 4.

3. TAYLOR, J. D., KENNEDY, W. J. & HALL, A. The Shell Structure of Mineralogy
at the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125; 29 Plates,
77 Text-figures. 1969. 4.50.

4. HAYNES, J. R. Cardigan Bay recent Foraminifera (Cruises of the R.V. Antur)
1962-1964. (In press.)

Printed in England by Staples Printers Limited at their Kettering, Northants, establishment

INDEX TO VOLUME 21

The page numbers of principal references and the new taxonomic names are printed in bold type

Abarenicola . . . . 108, in, 187

Ablepharus . . . . . .252

Acanthaster . . . . . 117

Acanthocaris . . . . 15

Acanthodactylus

234-236, 237, 249, 250, 251, 252, PI. 4
aceratus, Chaenocephalus . 53, 54, 64

acetabulosus, Tadarida .... 48

acetabulosus natalensis, Tadarida . . 48

Acropora 113, 131, 132, 133, 139, 148, 160,

161, 177, 179, 196, 198, 199, 202

acrotis, Rhinolophus clivosus
aculeifer, Hypoaspis
aeruginosa, Polispilota .
afer, Triaenops persicus .
affinis, Praxillella .
afra, Eunice .
africana, Lironeca
africana, Tadarida
africanus, Miniopterus inflatus

41,

PI. 3
. 188

107, 112, 157

. 329-338

Agama

215, 218, 222-232, 249,
250, 251, 252, Pis 1-3
222-232, 252
105, 107, 116, 154-155
. 163
223

106, 112, 114, 137
112, 117, 129
106, 134

359

- 359, 360

- 351-357
308, 309

106, 112, 113, 142

Agamidae
Aglaophamus
Aglaurides
agnetae, Agama
Agrius ....
albini, Synelmis
albolineatum, Phascolosoma
albopicta, Eulalia .
alcyone, Rhinolophus
alepidotus, Paraphoxinus
alepidotus, Phoxinellus .
Aliolimnatis
Allopolybothrus
alternata, Typosyllis
alticolus, Rhinolophus
alticolus, Rhinolophus simulator . . 37
amabilis, Malurus . 313, 314, 315, 317-318,

320, 322, 324, 325
Amaea

Amaeana . . .
amboinensis, Lycastopsis
amboinensis, Namanereis
americanus, Lithobius
Ammotrypane
Ampharetidae
Ampharetinae
Amphinome .
Amphinomidae
Amphiroa . . .
ampullifera, Harmothoe
ampullifera, Polynoe

195
108, 195

147
117, 118, 147

297, 299, 302, 309

. 182

105, 108, 115, 193-194

194
106, 113, 130

105, 106, 130-133

112, 126, 131, 151

126

ampulliferus, Paralepidonotus
Anaitides . . 106, in, 118,
ananas, Thelenota .
Ancistrosyllis

Angel, M. V. . . 257

angolensis, Hipposideros caffer
Angosoma ....
Angosomaspis
Anguidae ....
anomala, Pseudonereis .
antarctica, Ogmogaster .
antennata, Eunice . 107,

antillensis, Eunice .
antipoda, Pectinaria
anurus, Epomophorus
Aphroditidae . 104, 105,

aphroditois, Eunice
apodus, Ophisaurus
Arabella . . 107,

Arabellinae ....
arenaceodonta, Neanthes
arenaceodonta, Nereis
Arenicola ....
Arenicolidae ....
argus, Bohadschia .
argus, Lithobius

Armandia . . . 108,

armillaris, Syllis
armillaris, Typosyllis
Asellia .....
asper, Acanthodactylus boskianus
aspergillum, Cloeosiphon
aspersa, Lacerta
Aspidosiphon

assimilis, Malurus 313, 314, 315,
321, 322, 323, 324,
assimilis, Malurus lamberti

106, 126-127
"9, 135-136

117, 120

136, 137

-283, 289-296

38, 39, 4

92-94, 96, 97

253

107, 112, 151

54, 55
ii2, 114, 157

342

108, 116, 193

. 29

106, 119-129

107, 113, 157

253

112, 113, 164
164-165

. 148
. 148
. I8 7

105, 108, 187

117, 120

303, 309

113, 116, 182

. 142

106, 112, 142

40-41
234, 249, 252

112, 117, 129

234
112, 117, 129

316, 318, 320,
325, 326, 327

319

, 86, 96, 97
117, 120
117, 120
. 179
. 96

342

athiasae, Hypoaspis . 81-84

atra, Halodeima

atra, Holothuria

Audouinia ....

aurata, Cetonia

auriculata, Eunice

australiensis, Latreillia .

australis, Taphozous perforatus . . 33

Autolytinae ... 139

Autolytus 1 06, 139

Balaenoptera . . . -53, 54- 56

balaenopterae, Diplogonoporus 54, 56

Balanoglossus . . .113, 117, 125
bandaensis, Sigalion . . 106, 113, 128

364

INDEX

20
25-49

343

3, 9, 10, 13, 16, 18
313, 318, 320

barbata, Homola .
Bats ....
benedicti, Eunice .
benedicti, Raninoides
bernieri, Malurus .
bernieri, Malurus lamberti . . .319
Bhawania . . 106, 112, 117, 129-130

bifurcatus, Haploscoloplos 107, in, 116, 165
bimucronatus, Promacrus ... 96
bioculatum, Megalomma . . . 200

bisaccata, Hymenolepis ... 53, 60

bispinosus, Pentodon .... 74
blanfordi, Agama .... 229, 251
blanfordi, Ophisops .... 240
blanfordi, Ophisops elegans 238-241, 250, 252
blanfordi fieldi, Agama 228-230, 250, 252, PI. 3
blanfordii, Bunopus . . . .252
bocagei, Myotis ..... 44
Bohadschia . . . . . 117, 120
boskianus asper, Acanthodactylus 234, 249, 252
Bothriocephalus .... 54, 56-58
Bothropolys ..... 302, 309
brachyaskos, Conchoecia . . 281, 282

brachycola, Syllis . . . . .143
brachycola, Typosyllis

106, in, 112, 113, 114, 143

brachydactyla, Agama stellio . 231-232, 252
brachypedes, Lucanaspis
Branchiomma
Brania ....
brevicirrus, Perinereis nuntia
brevicornis, Lithobius
brevipes, Ophiocoma
brevirostris, Eremias

. 88-90, 97
108, 113, 198-199, 200
106, 139
. 150
299, 303-304, 309

117, I2O

237

brevirostris, Eremias brevirostris . .237
brevirostris brevirostris, Eremias . . 237
brevirostris microlepis, Eremias

236-237, 2 5 2 5 2

buchnerorum, Lithobius castaneus . 300, 308
Bunopus ...... 252

caducus, Dasybranchus .

caffer, Hipposideros

caffer, Hipposideros caffer

caffer angolensis, Hipposideros

caffer caffer, Hipposideros

caffer nanus, Hipposideros

caffer tephrus, Hipposideros .

cantoris schmidti, Acanthodactylus

capensis, Eptesicus

capensis, Gyptis

capensis, Mediomastus .

capensis, Ophelia .

capensis, Oxydromus

Capitellethus

Capitellidae .

Capitobranchus

Cardioderma

carnosus, Balanoglossus .

108, 113, 185

38, 39, 4
. 38-39, 40

38,39
38,39
252
42

106, 137, 138
108, 186

- 183

137
108, 184

105, 1 08, 184-186
108, 113, 184-185

35
113, 117, 125

105,

232-

Carobia ....

castanea, Carobia .
castanea, Genetyllis
castanea, Phyllodoce
castaneus, Lithobius
castaneus buchnerorum, Lithobius
caudata, Neanthes
caudata, Nereis
celerio, Hippotion .
centaurus, Angosoma
centralis, Hipposideros ruber . 38
Ceramodactylus
Cerastes ....
cerastes, Cerastes .
Ceratonereis . . 107, 113,

Cerithium ....
Cetonia ....

Chaenocephalus
Chaetopteridae
Chaetopterus
Chalcides
Chamaeleo
chamaeleon, Lacerta
chamaeleon musae, Chamaeleo
chamaeleon recticrista, Chamaeleo

232-

Chamaeleonidae
channeri, Lyreidus
chestnut-shouldered wren complex
Chilopoda ....
chimmonis, Notosceles .
chinensis, Leocrates
Chiroptera ....
Chloeia ....

Chloephaga ....
chloronotus, Stichopus .
Chrysopteron
Cichlid fishes

cingulata, Branchiomma 108,

cingulata, Dasychone
cirrata, Laonice

Cirratulidae . . . 105,

cirratulus, Nerine .
cirratulus saipanensis, Nerine .
cirrifera, Prionospio
Cirriformia . . . 107,

claparedii, Abarenicola .
claparedii, Abarenicola claparedii
claparedii, Arenicola
claparedii, Leocrates
claparedii, Sphaerodoridium .
claparedii claparedii, Abarenicola
clarus, Malurus
clavata, Brania
clavata, Grubea

clavigera, Gastrolepidia . 106, 116,
clivosus acrotis, Rhinolophus .
Cloeosiphon ....
Clymenella . . . 108,

coasta, Cossura

. 136

1 06, 136

106, 136
300, 308, 309

300, 308

107, 148
107, 148

, 39, 40, PI. 3

252

253

253
147-148, 153

117, 160
. 96
53, 54- 64
107, 177-179
107, 177
241, 249, 252
234, 249, 252
232

234

234, 249, 252
232-234, 252

4, 9
313-328
297-311

106, 113, 139
25- 4 9

106, 116, 131
53, 54, 58, 61

117, I2O

329-338

113, 198-199

. 198

107, 167

107, 179-180

174

J 74
107, 170

112, 113, 179
. 187

108, in, 187

. 187

139

107, in
108, in, 187

. 320

106, 139

139

117, 119-I2O

112, 117, 129

113, 187-188

108, 180

INDEX

365

coccinea, Eunice . 107, 112, 118, 119, 158

coeliaca, Trypanosyllis . 106, in, 112, 142

Coleolaelaps . . . 69, 76, 78, 88, 92, 96
collaris, Eirenis ..... 253
collaris, Lysidice . 107, 112, 114, 118, 119, 162
colorata, Echis ..... 253
Coluber . 215, 244-245, 249, 253, PI. 6

Colubridae .... 243-246, 253
commensalis, Hololepidella . . 121, 122
complanata, Eurythoe

106, 112, 113, 117, 130, 131

Conchoecia . . .
conspicua, Chloeia .

Contracaecum . .
convolvuli, Agrius ..
cor, Cardioderma ..
corallicola, Pseudopolydora
cornuta, Ehlersia . . .
cornuta, Langerhansia . .
cornuta, Syllis . .
Coronella . .
coronella, Eirenis coronella .
coronella coronella, Eirenis .
coronella fraseri, Eirenis .

corsicus, Lithobius impressus .
corsicus, Polybothrus impressus
Corynosoma . . . .
Coryphaena . . .
Cosmonotus . . . .
Cossura

Cossuridae . . .
Costa, M
costae, Ceratonereis .

257-283, 289-296
. 106, 116, 131
. . 54, 64
... 34
... 35
107, in, 116, 173
. . .141
. 106, 113, 141
.141
. 243

. . . 253
. . . 253
. . -253
. . . 307
. 307
54. 64

53. 54. 5^
. 4, 9, 13, 15
108, 180

105, 108, 115, 180
67-gS
. . 107, 147

costarum, Spiochaetopterus costarum

107, in, 113, 178-179
costarum costarum, Spiochaetopterus

107, in, 113, 178-179

costarum monroi, Spiochaetopterus . .179
costarum okudai, Spiochaetopterus . .179
costarum pottsi, Spiochaetopterus . .179
coutierei, Pomatoceropsis . . . 204
coutierei, Spirobranchus . 108, 113, 204

cricognatha, Nereis . . . .148

crinoidicola, Paradyte . 106, 116, 117, 126

crinoidicolo, Polynoe . . . .126
cristata, Pista . . . . 197

cryptocephala, Bhawania . . 129, 130
cryptocephala pottsiana, Bhawania . .130
Cryptonereis . 105, 107, 117, 118, 144-147
crypturus, Epomophorus . . .29

cultrifera, Perinereis 107, 113, 118, 119, 149

cumingi, Paraspidosiphon . . 117, 125
cyaneus, Malurus ..... 325
Cymothoidae .... 329-338

Cyprinidae ..... 359-361
cystops, Rhinopoma hardwickei . . 30

Dasybranchus
Dasychone
debilis, Paleanotus
decemlineata, Eirenis
decipiens, Conchoecia

dentata, Ophiocoma insularia
denti, Rhinolophus
denticulata, Ophelia
Diadema

diadema, Spalerosophis
diagramma, Simochromis
dianchora, Neofolitispa .
dibranchis, Aglaophamus
dibranchis, Nephtys
dibranchis, Pista

digitata, Thalenessa

Diogena

Diphyllobothrium .

Diplocirrus .

Diplogonoporus

'Discovery* expeditions, 1925-

dispar, Capitellethus

Dispio . . . 105,

diversa, Aliolimnatis .

dobsoni, Rhinolophus landeri

Dodecaceria .

dollfusi, Mastobranchus .

doriae, Ceramodactylus .

dorsipes, Notopus .

Dorveilleinae

Dorvillea

Drilonereis

dubius, Hypoaspis

dulcis, Malurus

Dynastaspis

Dynastes
dysteri, Salmacina

108, 113, 185

. 198

106, 112, 130

253
259, 269, 279, 280,

281, 282

117, 120

253

329
141, 142, 178

154

105, 108, 113, 114,

116, 196-197

1 06, 129

34
54. 55, 56

105, 108, 116, 181

54- 56

-1936 . 51-65

. ~ . 108, 184

107, 113, 166-167
. 351-35 7

1 08, 112, 179-180

108, in, 116, 185

252
4. 15

. 165

107, 165

. 84-86, 97

313, 314, 315, 317,

321, 322, 324, 325

90-92, 96, 97

204

danfordi, Lacerta danfordi
danfordi danfordi, Lacerta

252

Eason, E. H.
Echis . . .
eetweldii, Eugnathichthys
ehlersi, Euleanira .
ehlersi, Lygdamis .
ehlersi, Potamilla .
ehlersi, Prionospio
ehlersi, Sthenelanella
ehlersi, Tetreres
ehrenbergi, Ophisops elegans
239,

ehrenbergi, Terebella
Eirenis

elegans, Aspidosiphon
elegans, Lepidasthenia .
elegans, Malurus . 3 Z 3.
elegans, Ophiarthrum
elegans, Ophisops .

. 297-3H

253
329, 337

. 128
108, 192
108, 201

107, in, 116, 170

106, 116, 128-129

192

240, 241, 250, 252
108, 112, 198

253

112, 117, 129

106, 113, 117, 123
314, 315, 322, 323

117, 120
238, 240, 241

366

INDEX

elegans, Ophisops elegans . . 239, 252
elegans blanfordi, Ophisops 288-241, 250, 252
elegans ehrenbergi, Ophisops

239, 240, 241, 250, 252

elegans elegans, Ophisops . . 239, 252
elegans schlueteri, Ophisops . . 239, 241
elioti, Phyllochaetopterus 107, in, 113, 134, 177
elongatus, Eupolybothrus . . . 308
elongatus, Lamprologus . . . 329, 335
elongatus, Lithobius 299, 305-3O6, 307, 309

elongatus, Polybothrus .... 307
elongatus impressus, Eupolybothrus . 308

elongatus koenigi, Polybothrus . . 307
elongatus oraniensis, Lithobius . . 307
elongatus oraniensis, Polybothrus . . 307

emarginatus, Henicops
emarginatus, Lamyctes
emarginatus, Lithobius
enigmatica, Lironeca
enigmatica, Mercierella
Epomophorus
Eptesicus
Eremias

erythraeensis, Ceratonereis

eson, Hippotion

Eteone

Euchloron

Euclymene

Euclymeninae

Eugnathichthys

Eulalia

Euleanira

Eumeces

Eunice

Eunicidae

Eunicinae

Euphrosine .

Eupolybothrus

Eupolymnia .

Euprepes

Eurato

eurystoma, Myriochele

Eurythoe

Euthelepus

Euthynnus .

exilis, Syllis .

exilis, Typosyllis .

eximius, Lithobius

Exogone

Exogoninae .

expansus, Ichthyoxenus

expansus, Lironeca

297, 300
301
297, 300-301, 309

329, 335. 336, 337
108, 117, 203
29
42
236-238, 249, 250, 252

107, 147

34
106, in, 113, 133

108, 188
108, 187-189

329, 337
106, 112, 134

128

241-243, 249, 250, 252, 253, PI. 5
105, 107, 112, 113, 114, 116, 117,
118-119, 14. 157-162, 339-344
104, 105, 107, 157-165, 339-344
157-162
106, 112, 131
306, 308, 309
108, 114, 195, 198
. 241
202

108, 116, 189-igo
106, 112, 113, 117, 130, 131-132
108, 114, 195
53, 54. 55

143
107, 143

300
106, in, 112, 140

139-140
329, 337
329, 335, 337

fasciatus, Lithobius . . . 301, 309
fausta, Diogena ..... 34
fieldi, Agama blanfordi 228-230, 250, 252, PI. 3
fieldi, Agama persica . . . 228, 250

fieldi, Pseudocerastes .... 253

filicornis, Spio .... 107, 174

filigera, Audouinia . . . .179

filigera, Cirriformia . . 107, 113, 179

Filograna ..... 108, 204

Filograninae ...... 204

fistulicola, Dodecaceria . 108, 112, 179-i8o

Flabelligeridae . . 105, 108, 115, 181

flavipunctatus guweirensis, Pristurus . 252
floricola, Potosia ..... 96

floridana, Eunice . . . . .160

foliosa, Euphrosine

forficatus, Lithobius

fraseri, Eirenis coronella

fristedti, Phyllodoce

fulgida, Aglaurides

fulgida, Oenone

fullo, Polyphylla .

fumigatus, Rhinolophus fumigatus

fumigatus fumigatus, Rhinolophus

fusca, Phoebetria .

fusca, Sabella

106, 131
297. 298, 302, 303, 309

253
106, in, 112, 134

163

107, 163
78, 96

. 37
53, 54, 62, 63

108, 201

fuscatus, Pipistrellus kuhlii
fusiformis, Owenia

108, 116, 192

Galaxea . . . 132, 134, 143, 148

gallapagensis, Pseudonereis . . .151
Gamasus ...... 70

gambianus, Epomophorus ... 29
gangetica, Platanista . . . 345-348
Gastrolepidia . 106, 116, 117, 119-120

Gekkonidae .... 219-222, 252

gemmifera, Exogone . . 106, 112, 140
Genetyllis ..... 106, 136

Gibbs, P. E. . . . . 99-2H

gigantea, Glycera . . 107, 113, 116, 156

giganteus, Spirobranchus 108, 113, 203-2O4

gilchristi, Nerinides . . . 107, 168
glabratus, Lithobius .... 304

Glandicephalus .... 54, 55, 56

glandigerus, Vermiliopsis . 108, 112, 204

Glauconycteris .... 42-43

glaucus, Diplocirrus . . . .181

glaucus orientalis, Diplocirrus

105, 108, 116, 181

Glycera . . 107, 113, 116, 156-157, 205

Glyceridae . . 105, 107, 156-157, 205

Glycerinae ..... 156-157

goodei, Bhawania . 1 06, 112, 117, 129, 130

gracilipes, Lithobius . . . .301

gracilis, Genetyllis . ... 106, 136

gracilis, Phyllodoce . . . 106, 136

gracilis, Syllis . . . . 1 06, 142

graft ei, Bohadschia . . . 117, 120

grandiceps, Stenodactylus . . 216, 252
grandis, Acanthodactylus . 235-236, 250,

252, PI. 4
grayi, Cosmonotus ... . 4

Great Abbai Expedition, 1968 . 25-49

INDEX

367

gregoryi, Notopygos . 106, in, 119, 132

griseus, Varanus griseus .... 253

griseus griseus, Varanus .... 253

grossipes, Lithobius . . . .301

Grubea ...... 139

grubei, Eunice . . . 107, 114, 158
guadalcanalis, Lepidasthenia

105, 106, 113, 117, 123-125

guineensis, Hipposideros ruber . 38, 40

guttata, Ptyodactylus hasselquistii . .220
guttatus, Ptyodactylus . . . .220

guttatus, Ptyodactylus hasselquistii

220-221, 250, 252

guttulata, Eremias . . . -237

guttulata, Eremias guttulata 237-238, 249, 252
guttulata, Lacerta . . . .237

guttulata guttulata, Eremias 237-238, 249, 252
guttulata watsonana, Eremias . . 252

guweirensis, Pristurus flavipunctatus . 252
Gyanisa ...... 34

Gyptis . . 105, 106, 114, 137-138

haasi, Agama pallida

215, 222-228, 250, 252, PI. i
haedinus, Taphozous perforatus . 31-34

Haemopis ...... 350

Halichondria . 130, 139, 140, 141, 142, 143,

144, 147, 148, 162, 177, 183

Halimeda
Halodeima .
hamanni, Corynosoma
hanaokai, Ancistrosyllis
hanaokai, Sigambra
Haploscoloplos
Haplosyllis .
hardwickei, Lithobius
hardwickei, Rhinopoma

163, 196
117, 120
54,64
. 136
116, 136

106, in,

107, in, 116, 165

106, 141

297-298, 299, 309
30

hardwickei cystops, Rhinopoma . . 30
hardwickei macinnesi, Rhinopoma . . 30
hardwickei sennaarinese, Rhinopoma

30-31, PL 3

hardyi, Acanthodactylus scutellatus . 251

Harmothoe ..... 106, 120

Harrison, C. J. 313-328

hartmanae, Serpula . . 108, in, 203
hasselquistii, Ptyodactylus . . 221, 222
hasselquistii, Ptyodactylus hasselquistii . 250
hasselquistii guttata, Ptyodactylus . .220
hasselquistii guttatus, Ptyodactylus

220-221, 250, 252

hasselquistii hasselquistii, Ptyodactylus . 250
hasselquistii puiseuxi, Ptyodactylus

222, 250, 252

Hemidactylus . . 219-220, 249, 252

hendersoni, Raninoides . . . . 4, 18
Henicops . . . . 297, 300, 301
herdmani, Phyllochaetopterus

107, in, 113, 117, 125, 178
heruensis, Myriochele 105, 108, 116, 190-I9I

Hesione

Hesionidae .

heterochela, Sthenelais .

heteroclitus, Tetrabothrius

Heteroligus .

Hill, J. E. .

Himerometra

Hipposideros

hipposideros minimus, Rhinolophus

Hippotion

hirsuta, Sphaerosyllis

Hirudinaria .

Hirudinea

Hirudinidae, S. L. .

Hirudo

holobranchiata, Nothria

holobranchiata, Onuphis

Hololepidella

Holothuria

Homola

Homolidae

106, 114, 139
105, 106, 137-139

1 06, 1 1 6, 128

54, 62-63

25-49
117, 126
33, 38-40, PI. 3
35-36

34
106, 140

353
349-357
349-357

349

107, 113, 116
107, 113, 116, 163

105, 106, 114, 116, 117, 120-123
117, 120
18-22
18-24

honiarae, Poecilochaetus serpens

105, 107, 113, 175-176

hunteri, Promacrolaelaps . . 94-96, 97
Hyboscolex .... 108, 113, 181
Hydroides . . . . 108, 112, 202
Hydrurga . . . . 53, 54, 64

Hymenolepis . . . 53, 54, 58-61

Hypoaspis ..... 67-g8

Hypsicomus . . . 108, 112, 113, 199

Ichthyoxenus . . . . -329

impar, Harmothoe . . . .120

impatiens, Lumbrioconereis . . . 343
implexa, Filograna . . . 108, 204

impressus, Eupolybothrus . 306, 308, 309
impressus, Eupolybothrus impressus . 308

impressus, Lithobius . . 305, 306, 307
impressus corsicus, Lithobius . . . 307
impressus corsicus, Polybothrus . . 307
impressus elongatus, Eupolybothrus . 308

impressus impressus, Eupolybothrus . 308

indica, Lagisca . . . . .127
indica, Lycastis . . . . .147
indica, Namalycastis . . 107, 117, 147
indica, Sabellastarte .... 202
indica, Scolelepis . . . . .168
indicus, Lygdamis . . . 108, 192, 193
indicus, Malacoceros . . 107, 113, 168
indicus, Paralepidonotus . . 106, 127

inflatum, Scalibregma . . . 108, 181
inflatus africanus, Miniopterus . . 46

insignis, Lithobius . . . 301, 309

insignitus, Coluber .... 245

insignitus, Malpolon monspessulanus

245-246, 249, 253

insolita, Platynereis . 107, in, 112, 150

insularia dentata, Ophiocoma . . 117, 120

3 68 INDEX

integer, Coleolaelaps .... 76

lamberti mastersi, Malurus

319

integer, Hypoaspis . 70, 71, 76-78, 96, 97

Lamprologus

329. 335, 336

integer, Laelaps . . . . ' . 76

Lamyctes ....

3 01 . 39

intermedium, Megalomma . . 108, 113

lancadivae, Glycera . 107,

113, 116, 156

International Indian Ocean Expedition . 1-24

lanceolata, Armandia . 108,

113, 116, 182

Iphione . . . 106, 113, 114, 123

landeri dobsoni, Rhinolophus .

. 35

iracensis, Acanthodactylus tristrami . 236

Langerhansia

106, 113, 141

iricolor, Arabella . . . 107, 113, 164

Laonice ....

107, 167

Isolda . . . . . 108, 116, 193

lashleyi, Diphyllobothrium

54- 55

isolepis, Agama . . . 229

latastii, Ophiomorus

253

Isopoda . . 329-338

latreilli, Lumbriconereis

163

latreilli, Lumbrineris . 107,

114, 163-I&4

Latreillia ....

20, 21

janickii, Bothriocephalus . . 54, 86-58

leachi, Lithobius .

. 298, 309

japanensis, Eteone . 106, in, 113, 133

Leachii, Lithobius .

. 298

japonica, Leanira . . . . .129

Leanira ....

129

japonica, Magelona . . 107, 113, 175

Leocrates ....

106, 113, 139

japonica, Paromola .... 20

Lepidasthenia 105, 106, in, 113,

117, 123-126

japonica, Sthenolepis . . 106, 116, 129

Lepidonotus . . . 106,

112, 113, 126

johnsoni, Poecilochaetus . . .176

leptocirrus, Armandia

108, 116, 182

jugularis, Coluber jugularis . . . 253

leptocirrus, Ophelina

. 182

jugularis jugularis, Coluber . . . 253

Leptonychotes . . 53,

54- 55. 56, 64

jukesi, Lepidonotus . 1 06, 112, 113, 126

leptonyx, Hydrurga

53. 54. 64

jukesi, Thormora . . 106, 112, 113, 126

Leptotyphlopidae

253

Leptotyphlops

253

Leuciscus ....

360

kerguelensis, Neanthes . . . 107, 148

leucoptera, Chloephaga picta .

53. 54. 58, 61

kerguelensis, Nereis . . . 107, 148

leucopterus, Malurus

327

kinsemboensis, Euthelepus . 108, 114, 195

Limnatis ....

349-357

kitaibelii, Ablepharus . . . .252

Limnobdella ....

349

koehleri, Macrophiothrix . . 117, 123

Limnothrissa

329, 337

koenigi, Lithobius ..... 305

linaresi, Megalomma . 108,

in, 113, 199

koenigi, Polybothrus .... 305

Lincoln, R. J.

. 329-338

koenigi, Polybothrus elongatus . . 307

Lironeca ....

329-338

kollikeri, Praxilla 188

Lithobiomorpha

297-311

kollikeri, Praxillella . . . .188

Lithobius ....

. 297-3H

koloana, Ophelia . 105, 108, 116, 182-i83

Lithozoea ....

15. 23

krameri, Gamasus ..... 70

lobatus syriacus, Ptyodactylus

222

krameri, Hypoaspis . . 70, 81, 84, 96

Loimia . . . 108,

113, 116, 196

kuhlii fuscatus, Pipistrellus ... 42

Longbottom, M. R. . ".

339-3 4 4

longicaudata, Armandia

108, 116, 182

longicirrus, Scalisetosus .

. 126

labiata, Nycteris thebaica . . 34-35

longipinnis, Glycera

107, 156

labiatus, Epomophorus .... 29

longiseta, Hyboscolex

108, 113, 181

Lacerta . .219, 232, 234, 237, 241, 252

longissima, Syllis . . 106,

III, 112, 141

Lacertidae .... 234-241, 252

Lucanaspis ....

88-90, 96, 97

Lacertilia ..... 219-243

Lucanus ....

70, 96

Lacydoniidae . 105, 107, 115, 155-156

lucasi, Lithobius

308, 309

ladacensis, Coluber rhodorhachis . . 244

lucida, Pionosyllis .

143

ladacensis, Zamenis .... 244

lucida, Typosyllis .

107, in, 143

laddi, Dodecaceria . . . 108, 180

Lumbriconereis

343

Laelaps ...... 76

Lumbrinereis

343

laevigata, Naineris . . 107, 113, 166

Lumbrinerinae

163-164

Lagisca ...... 127

Lumbrineris . 107, in, 114, 116,

119, 163-164

lamberti, Malurus . . 313, 314, 315, 316,

Lycastis ....

3i8, 319, 322, 324

Lycastopsis ....

- M7

lamberti, Malurus lamberti . . -319

Lycoris ....

. 151

lamberti assimilis, Malurus . . -319

Lygdamis ....

108, 192-193

lamberti bernieri, Malurus . . -319

Lyreidus . . 3, 4, 7,

9, 13, 15. 23

lamberti lamberti, Malurus . . .319

lyrochaeta, Aglaophamus

155

INDEX

369

lyrochaeta, Nephtys . . . 155

Lysaretinae . . . . . .163

Lysidice . . 107, 112, 114, 118, 119, 162
Lysilla .... 108, 113, 116, 195

Mabuia .....

Mabuya .... 243,
macgregori, Capitobranchus
macinnesi, Rhinopoma hardwickei .
macintosh!, Marphysa . . 107,
Macroclymene . . . 108,
Macrophiothrix ....
macrophthalma, Nudaurelea .
Macrophyllum ....
macroprocera, Conchoecia

259, 269,
278, 280,
106, 113,
106, 118,
106, 114, 118,
1 06,

maculata, Lepidasthenia
madeirensis, Anaitides
madeirensis, Phyllodoce
magalhaensis, Eulalia
magalhaensis, Pterocirrus . 106,
Magelona .... 107,
Magelonidae . . . . 105,
magna, Dispio ....
magna group ....

maja, Guyanisa ....
major, Drilonereis ....
majusculus, Triaenops persicus
Malacoceros .... 107,
malaitae, Gryptonereis

105, 107, 117, 118,

malayensis, Prionospio steenstrupi 107,
Maldanidae . . . 105, 108,

malmgreni, Phyllodoce . . 106,
malmgreni, Prionospio . . 107,
Malpolon . . 215, 245-246,

Malurus .....
manihine, Eunice
marauensis, Solomononereis

105, 107, 151,

maraunibinae, Gyptis 105, 106, 114,
maritimus, Taphozous
Markowski, S.

maroroi, Dispio . 105, 107, 113,

marovoi, Eunice . 105, 107, 116, 117,
Marphysa .... 107,
martiensseni, Otomops martiensseni
martiensseni martiensseni, Otomops
masalacensis, Lycoris
masalacensis, Nereis
masalacensis, Perinereis .
masalacensis, Pseudonereis 107, in,
mastersi, Malurus . . . 313,
mastersi, Malurus lamberti
Mastobranchus . 108, in, 113, 116,
maxillosus, Polyodontes . 106,

Mediomastus ....

medusa, Loimia . . 108, 113,

megabranchia, Eunice

243
249, 253

. 185
30

in, 162
"3, 187
117. 123

134
270-275,
281, 282
117, 125
119, 135
119, 135
112, 134

112, 134
"3. 175
197. 175

167
289, 291

107, 165
41

113, 168

144-147
116, 171
187-189
113, 135
170-171
249, 253
313-328
339-344

152-153
137-138

3 1

51-65
166-167

158-i6o

in, 162

. 46

. 119

112, 151
318, 320

319
185-186

113, 127
108, 186
1 1 6, 196

343

megaera, Euchloron .... 34

Megalomma . . 108, in, 113, 116, 199-201
melanocephalus, Rhynchocalamus . . 253
melanonotus, Polyodontes . . 106, 127
melanops, Lithobius . . . 304, 309
melanostigma, Sabella . . 108, 201-2O2
meles, Heteroligus .... 88

Melinninae ...... 193

mendanai, Scolelepis squamata

105, 107, 113, 174

Mercierella .... 108, 117, 203
Mesochaetopterus . 107, 113, 117, 123, 177

Mesostigmata .... 67-98

michaelseni, Lepidasthenia . . -125
microlepis, Eremias brevirostris

236-237, 250, 252
microlepis, Lepidasthenia
Micronephthys
Micronereides
Micronereis .
microphyllum, Rhinopoma
microprocera, Conchoecia

106, 113, 117, 125
107, 155
146
146

259, 264-27O, 272,
274, 278, 280, 282
30

Micropteropus ....

Millepora . . . . . .178

minax, Hydroides . . . 108, 112, 202
minax, Serpula ..... 202

minimus, Rhinolophus hipposideros 35-36

Miniopterus ...... 46

minor, Myriochele ..... 192

minuta, Hololepidella . . . 120, 121
minuta, Polynoe . . . .121

minuta oculata, Polynoe . .121

minutus, Mesochaetopterus . . .177
miodon, Limnothrissa .... 329

mirabilis, Ceratonereis . . 107, 113, 148
mirasetis, Aglaophamus . . 155

mirasetis, Nephtys 155

misakiensis, Notopus . 15

moilensis, Coluber . 245

moilensis, Malpolon . 215, 245, 249, 253
monilicornis, Lithobius 299, 305, 306-308, 309
monoceros, Hydroides . 202

monoceros, Oryctes . 69, 84, 86
monroi, Spiochaetopterus costarum . 179

monspessulanus insignitus, Malpolon

245-246, 249, 253

Morris, P. . . 25 ~49

morrisi, Chrysopteron . 44

morrisi, Myotis . 43-46, Pis. 1-2

Morton, J. E. .205

mossambica, Lepidasthenia . - 123

multidentatus, Bothropolys . 302, 303, 309
multidentatus, Lithobius . 302, 303, 309

multifilis, Tharyx . .180

multisetosus, Angosomaspis . 92-94, 97
munamaorii, Aglaophamus

105, 107, 116, 154-155

munamaorii, Nephtys 105, 107, 116, 154-155
muricata, Iphione . . 106, 113, 114, 123

INDEX

musae, Chamaeleo chamaeleon

musculus, Balaenoptera .

mutabilis, Agama .

mutans, Arabella .

Myotis .

Myriochele

myrtosa, Euphrosine

234
53, 54, 56

. 228
107, 112, 164

43-46, PlS. 1-2

105, 108, 116, 189-192
106, 112, 131

Naineris . . . 107, 113, 166

najadum, Coluber ..... 253
Namalycastis . . . 107, 117, 147

Namanereinae .... 144-147
Namanereis . . . 117, 118, 146, 147

nanus, Hipposideros caffer . 38, 39

nanus, Pipistrellus .... 42

nasicornis, Oryctes . . 70,71, 78, 96

natalensis, Tadarida acetabulosus . . 48

Natrix 243, 249, 253

Neanthes 107, 148

nebulosa, Eupolymnia . . 108, 114, 195
nebulosa, Polymnia . . . 195

neglecta, Ophelia . . . . .183

Nematonereis . . 107, 112, 113, 162

neocaledonica, Perinereis . . . 205

Neofolitispa . . . . 141, 142, 178

neokrameri, Hypoaspis 71, 72, 74, 96, 97

Neopomatus . . . . .117, 203

Nephele ...... 34

Nephtyidae . . 105, 107, 115, 154-155

Nephtys . . 105, 107, 116, 154-155

Nereidae . . 105, 107, 144-153, 205

Nereinae ..... 147-153

Nereis .... 107, 148, 150, 151

Nerine ....... 174

Nerinides 107, 168

Newport, G ,. 297-311

niapu, Hipposideros ruber . . 38, 40

PNicomache .... 108, 113, 189
Nicomachinae . . . . .189
nigeriae, Tadarida nigeriae . . 47, PL 3
nigeriae nigeriae, Tadarida . . 47, PI. 3
nigricans, Harmothoe . . . 106, 120
nigriceps, Tarbophis . . . .253
nigrobranchiata, Amphinome . 106, 113, 130
nigropunctata, Hololepidella

I06, 114, 117, 120-121, 122

nigro-punctata, Nereis . . . .150

nigropunctata, Perinereis 107, 112, 114, 150

nigropunctata, Polyeunoa . . .120

nigro-punctata, Polynoe . . .120
nilotica, Limnatis .... 349-357

Nipa . . . . . . .117

nobilis, Bothropolys .... 302

norvegica, Eunice . . 107, 160-i6i, 343

Nothria . . . 107, 113, 116, 163

Notocirrus . . . . . .164

Notomastus . . . . . 108, 186

Notophyllum . . . 106, 114, 134

Notopus ... . . 4, 9, 13, 15

Notopygos . . 106, in, 113, 114, 118, 132
Notosceles . . . . 4, 9, J 3> 15

Notothenia ..... 53, 54, 64

novaegeorgiae, Polydorella

105, 107, 116, 168-170
Nudaurelea ...... 34

nudicornis, Lithobius .... 305

nummifer, Coluber ravergieri . . . 253
nuntia, Perinereis . . 107, 113, 149, 150

nuntia brevicirrus, Perinereis . . .150
Nycteris . . . . . 33, 34

oblonga, Conchoecia . . . 259, 264
ocellata, Lacerta . . . . .241

ocellatus, Chalcides . . . .241

ocellatus, Chalcides ocellatus . 241, 249, 252
ocellatus ocellatus, Chalcides . 241, 249, 252
octacantha, Hymenolepis ... 60
oculata, Polynoe minuta . . .121

oculifera, Pseudeurythoe . . 133

Oenone ..... 107, 163

Ogmogaster . 54, 55

okudai, Spiochaetopterus castarum . .179
olivieri schmidti, Eremias . . .252
Onuphinae . . . . . .163

Onuphis . . . 107, 113, 116, 163

Ophelia . . . 105, 108, 116, 182-183

Opheliidae . . . 105, 108, 182-184

Ophelina . . . . . .182

Ophiarthrum . . . 117, 120, 123

Ophidia ..... 243-246

Ophiocoma . . . . . 117, 120

Ophiomorus . . . . . -253

Ophisaurus . . . . . -253

Ophisops . . . 238-241, 250, 252

ophiuricola, Hololepidella

105, 106, 114, 117, 121-123

oraniensis, Lithobius elongatus . . 307
oraniensis, Polybothrus elongatus . . 307
Orbiniidae . . . 105, 107, 165-166

orientalis, Acanthodactylus tristrami . 236
orientalis, Diplocirrus glaucus

105, 108, 116, 181
orientalis, Homola .... 20

Oryctes . 69, 70, 71, 78, 81, 84, 86, 96

osculatum, Contracaecum . . 54, 64

osiris, Hippotion . . . . .34

Otomops ...... 46

ovalis, Notopus . . . . .15

Owenia .... 108, 116, 192

Oweniidae . . 105, 108, 115, 189-192

Oxydromus ...... 137

Pachycephala
pacificum, Megalomma
palatii, Nephthys .
palauensis, Eunice
Paleanotus .

326
201

107, 155

342
106, 112, 130

INDEX

pallida, Agama 218, 224, 225, 226, 251, Pis. 1-2
pallida, Agama pallida 223, 228, 250, Pis. 1-2
pallida haasi, Agama . . 215, 222-228,

250, 252, PI. i

pallida pallida, Agama . 223, 228, 250, Pis. 1-2
Palmyridae . . . 105, 106, 129-130

Palola . . 107, 112, 113, 118-119, 161-162
paluda, Limnatis ..... 350

pantherina, Hesione . . . 139

papillifera, Lumbriconereis . . .164
papillifera, Lumbrineris . 107, in, 116, 164

Paradyte . . . 106, 116, 117, 126

Paralacydonia . . 107, 116, 155-156

Paralepidonotus . . . 106, 126-127
Paraphoxinus .... 359-36 1

Pararhodeus ...... 360

Paraspidosiphon . . . .117, 125

Pareurythoe . . 106, in, 113, 114, 132

Paromola ..... 20, 21

parthenoda, Conchoecia . 289, 293, 294, 295
parva, Anaitides . . . 106, in, 136
parva, Phyllodoce . . . 106, in, 136
patrizii, Asellia ..... 41

paucibranchiata, Pseudeurythoe

106, 113, 116, 133

pavimentatus, Eumeces schneideri . 242, 250
Pectinaria .... 108, 116, 193
Pectinariidae . . 105, 1 08, 115, 193

pectoralis, Pachycephala . . .326

pelamis, Euthynnus . . -53, 54, 55
peneus, Nephele ..... 34
pennata, Eunice . . . . -343
Pentodon . . . . . 74, 96

pentodoni, Hypoaspis . . 71-74, 9 6 , 97
perfoliatus, Glandicephalus . . 54, 55, 56
perforatus, Taphozous . 31, 32, 33, 34, 40

perforatus, Taphozous perforatus . 31, 32, 33
perforatus australis, Taphozous . . 33
perforatus haedinus, Taphozous . 31-34

perforatus perforatus, Taphozous . 31, 32, 33
perforatus rhodesiae, Taphozous . 32, 33

perforatus Sudani, Taphozous . . 31, 32, 33
perforatus swirae, Taphozous . . 32, 33

Perinereis . . 107, 112, 113, 114, 118,

119, 149-150, 205
Perolepis . . . . . .126

persica, Agama .... 229, 230

persica fieldi, Agama . . . 228, 250
persicus afer, Triaenops . . . 41, PI. 3
persicus majusculus, Triaenops . . 41
personatus, Raninoides . . . . 4, n

peruana, Prionospio . . . .173

phaeotaenia, Hypsicomus 108, 112, 113, 199
phalaena, Glauconycteris variegata . . 42
phalangium, Latreillia .... 20

Phascolosoma . . . 112, 117, 129
philactes, Hymenolepis .... 60

phillipsi, Leptotyphlops . . . -253
Phoebetria . . . -53, 54, 62, 63
Phoxinellus 359-36 1

Phyllochaetopterus 107, in, 112, 113, 117,

125, 126, 129, 130, 131, 134,

135, 137. HO, ML 142, M3,
147, 150, 151, 157, 161, 162,

164, 177-178, 2OO, 2OI, 202

Phyllodoce . . 106, in, 112, 113, 114,

118, 119, 134-136

Phyllodocidae . . 105, 106, 133-136

phyllognathi, Hypoaspis 70, 74-76, 86, 96, 97
Phyllognathus ..... 76
picea, Agama stellio . . . .252
picta leucoptera, Chloephaga . 53, 54, 58, 61
pictum, Ophiarthrum . . .117, 123
pictus, Polyophthalmus . . 108, 183-184
Pilargidae . . . 105, 106, 136-137

pilicornis, Lithobius . 298-299, 3, 39

Filled, G 345-348

pinnata, Prionospio . . 107, 116, 171
Pionosyllis . . . . . .143

Pipistrellus . . . . . .42

Pisces 359-361

Pista . . 105, 108, 113, 114, 116, 196-198
pitipanaensis, Pareurythoe

106, in, 113, 114, 132

planci, Acanthaster . . . .117
planus, Lithobius . . 302-303, 309

Platanista 345~34 8

Platynereis . . . 107, in, 112, 150

platypus, Lithobius . . . 304, 309
Platysphinx . . . . . .34

pleurostigma, Lamprologus . . 329, 335
Poecilobdella . . 353

Poecilochaetus . 105, 107, in, 113, 175-176
Polispilota .... -34

Polybothrus . . . 35. 37

Polychaeta . . 99-2 n, 339-344

Polycirrinae . . . 195

Polydorella . . 105, 107, 116, 168-170

Polymnia ...... 195

Polynoe .... 120, 121, 126

Polynoinae ..... 119-127

Polyodontes . . . 106, 113, 127

Polyodontidae . . .127

Polyophthalmus . . . 108, 183-184
Polyphylla . . 7 8 . 96

Pomatoceropsis . . . 204

Porites . . 125, 130, 132, 133, 148,

158, 163, 179, 180,
199, 202, 203, 204

Potamilla 108, 201

Potamobdella 349

Potosia .... -96

pottsi, Spiochaetopterus costarum . 179

pottsiana, Bhawania . . 106, 117, 130
pottsiana, Bhawania cryptocephala . .130

Praxilla 188

Praxillella . . - 108, 188

princeps, Eumeces . . . 241, 243

princeps, Eumeces schneideri

241-242, 250, 252, PI. 5

372

INDEX

princeps, Euprepes . . . . 241

Prionospio . . 105, 107, in, 116, 170-173
Pristurus ...... 252

procera, Conchoecia 259-264, 266, 268, 269,

272, 274, 278, 280, 282

'procera' group .... 257-283

prolifera, Polydorella . . . 169, 170
prolifera, Syllis . . . . .143

prolifera, Typosyllis . . 107, 112, 143
Promacrolaelaps . . . 94-96, 97

Promacrus ...... 96

propitia, Lithobius . . . .304

proximus, Coleolaelaps .... 88

prudhoei, Hymenolepis . -54, 58-6 1

pruvoti, Phyllodoce . 106, in, 112, 135

Psammolyce . . . 106, in, 113, 128

Psammophis . . . . -253

Pseudeurythoe . . 106, 113, 116, 133

Pseudocerastes . . . . -253

Pseudonereis . . 107, in, 112, 151

pseudoparthenoda, Conchoecia . 289-2g6
Pseudophoxinus .... 359-36 1

Pseudopolydora . 107, in, 116, 117, 173-174
Pterocirrus .... 106, 112, 134

Ptyodactylus . . 220-222, 250, 252

puiseuxi, Ptyodactylus .... 222

puiseuxi, Ptyodactylus hasselquistii

222, 250, 252

pulchella, Isolda . . . 108, 116, 193
pulcherrimus, Malurus . 313, 314, 315-316,

322, 323, 324

pulchricornis, Lithobius . . . 298, 299
pumila, Tadarida .... 46-47

punctata, Cirriformia . . 107, 112, 179
punctatus, Pentodon .... 96

pusillus, Micropteropus .... 30

quadraticeps, Phyllodoce . 106, 112, 135

quadratum, Diphyllobothrium . 54, 56

quadrioculatum, Branchiomma . . 200
quadrioculatum, Megalomma . 108, in, 200
Quantenobdella ..... 349
queenslandia, Reteterebella 108, in, 113, 198

Ranina . 3, 4, 5, 6, 7, 9, 13, 15, 23

ranina, Ranina . . 3, 4, 5, 6, 7, 9, 13

Raninidae . . . . . . l-i8

Raninoides . 3, 4, 7-11, 13, 15, 16-18, 23

ravergieri nummifer. Coluber . . -253
recticrista, Chamaeleo chamaeleon

232-234, 249, 252

recticrista, Chamaelo vulgaris . . . 232
regularis, Perolepis . . . .126

remanei, Dispio . . . . .167

remilleti, Hypoaspis . . 70, 86-88, 97
reniformis, Potamilla . . . .201

Reteterebella . . 108, in, 113, 198

rhinoceros, Oryctes . . . 69, 8 1

rhinocerotis, Coleolaelaps . . 69, 78

rhinocerotis, Hypoaspis . 78-8 1, 84, 86, 96, 97
Rhinolophus .... 35~37

Rhinopoma .... 30-31, PI. 3

rhodesiae, Taphozous perforatus . 32, 33

rhodorhachis, Coluber . 215, 244, 249, 253
rhodorhachis, Coluber rhodorhachis 244, PL 6
rhodorhachis, Zamenis .... 244

rhodorhachis ladacensis, Coluber . . 244
rhodorhachis rhodorhachis, Coluber 244, PI. 6
rhodorhachis subnigra, Coluber . .244
Rhynchocalamus ..... 253

Rice, A. L. . . . . . . 1-24

rigida, Ancistrosyllis . . . .137

robustipinna, Himerometra . . 117, 126
robustus, Acanthodactylus . . . 252
rogersi, Coluber . 244-245, 249, 253, PI. 6
rogersi, Malurus . 313, 314, 315, 317, 318,

320, 321, 322, 324, 325
rogersi, Zamenis . . . . .244

Prosea, Langerhansia . . . 106, 141
rossi, Notothenia . . . -53, 54, 64
rothi, Eirenis ..... 253

rouxi, Glycera .... 107, 157

Roxburgh, W 345~34 8

ruber, Hipposideros . . 33, 38, 39, 40
ruber, Hipposideros ruber . . 38, 40

ruber centralis, Hipposideros . 38, 39, 40, PI. 3
ruber guineensis, Hipposideros . 38, 40

ruber niapu, Hipposideros . . 38, 40

ruber ruber, Hipposideros . . 38, 40

rubriceps, Lithobius . . . 301, 309
ruderata, Agama . . . . .228
Rutilus 360

Sabella . . . . 108, 201-202

Sabellariidae . . 105, 108, 192-193

Sabellastarte . . . 108, 113, 202

Sabellidae . . . 105, 108, 198-202

Sagittarius, Mesochaetopterus

107, 113, 117, 123, 177

saipanensis, Nerine cirratulus . 174

Salmacina ...... 204

sancti-josephi, Eurato .... 202

sanctijosephi, Sabellastarte . 108, 113, 202
sanguisuga, Haemopis .... 350

savignii, Agama . 250

savignyi, Diadema . . .117

Scalibregma 108,181

Scalibregmidae . . . 105, 108, 181
Scalisetosus . . . . . .126

schlueteri, Ophisops elegans . . 239, 241
schmidti, Acanthodactylus cantoris . . 252
schmidti, Eremias olivieri . . .252
schneideri, Eumeces . . 242, 250, PI. 5
schneideri, Eumeces schneideri

242-243, 249, 253, PI. 5
schneideri, Scincus .... 242

schneideri pavimentatus, Eumeces . 242, 250

INDEX

373

schneideri princeps, Eumeces

241-242, 250, 252, PI. 5
schneideri schneideri, Eumeces

242-243, 249, 253, PI. 5
schokari, Psammophis .... 253

schusterae, Dispio . . . . .167

Scincidae . . . 241-243, 252-253

Scincus . . 242, 243, 253

Scolelepis . . 105, 107, 113, 168, 174

Scotophilus ...... 46

scutata, Sternaspis . . . 108, 184

scutellatus, Acanthodactylus scutellatus

235. 236, 251

scutellatus hardyi, Acanthodactylus . 251

scutellatus scutellatus, Acanthodactylus

235, 236, 251
Selysius ...... 44

semicincta, Audouinia . . . .179

semisegregata, Eunice .... 343

senegalensis, Taphozous 33

sennaarinese, Rhinopoma hardwickei 30-31, PI. 3
sepsoides, Sphenops . . . 251, 253
serpens, Poecilochaetus . . . .176

serpens honiarae, Poecilochaetus

105, 107, 113, 175-176

Serpula . . . 108, in, 202, 203

Serpulidae . . . 105, 108, 202-204

Serpulinae ..... 202-204

serratifrons, Raninoides . . . 4, n, 18
serrulata, Lithozoea ... 15, 23

sibogae, Isolda ..... 193

sibogae, Lepidasthenia . . . .126

sibogae, Notopygos . 106, 113, 114, 132
siciliensis, Eunice . . . 107, 112, 113,

118-U9, 161-162

siciliensis, Palola . . . 107, 112, 113,

118-119, 161-162
Sigalion .... 106, 113, 128

Sigalioninae ..... 128-129

Sigambra . . . 106, in, 116, 136

silenus, Phyllognathus .... 76

Simochromis ..... 329

simulator, Rhinolophus . . . 36, 37

simulator, Rhinolophus simulator . . 37
simulator alticolus, Rhinolophus . . 37
simulator simulator, Rhinolophus . . 37
sinaita, Agama . 229, 230-231, 249, 252, PI. 3
sinaitus, Hemidactylus .... 220

sinensis, Aglaophamus . . . 155

sinensis, Nephyts . . . . 155

Siphonosoma . . 113, 117, 125, 126

Sittella 326

sloanei, Lithobius .... 300, 309

socialis, Phyllochaetopterus 107, 112, 126, 129,

130, 131, 135, 137, 140, 141, 142, 143, 147, 150,

I 5 I > I 57 J 6i, l62 , 164, 178, 200, 201, 202

Solomononereis . . 105, 107, 151-153

somalicus, Eptesicus somalicus . . 42

somalicus somalicus, Eptesicus . . 42

PSosane .... 108, 116, 194

Spalerosophis . -253

sphaerocephala, Lumbriconereis . .164
sphaerocephala, Lumbrineris . 107, 119, 164
sphaerocephalus, Notocirrus . . .164
sphaerocirrata, Micronephthys . 107, 155

sphaerocirrata, Nephthys . . 107, 155
Sphaerodoridae . . . 105, 107, 144
Sphaerodoridium .... 107, in

Sphaerosyllis .... 106, 140

Sphenops . .251, 253

Spio ...... 107, 174

Spiochaetopterus . 107, in, 113, 178-179

Spionidae . . . 105, 107, 166-174

Spirobranchus . . 108, 113, 203-204

splendens, Macrophyllum . . .134
splendens, Notophyllum . 106, 114, 134

splendida, Hesione . . 106, 114, 139

spongicola, Haplosyllis . . . 106, 141
spongicola, Syllis . . . . .141

squamata mendanai, Scolelepis

105, 107, 113, 174
steenstrupi, Prionospio . . . .171

steenstrupi malayensis, Prionospio 107, 116, 171
stellio, Agama .... 232, 249

stellio, Agama stellio .... 252

stellio brachydactyla, Agama . 231-232, 252
stellio picea, Agama .... 252

stellio stellio, Agama . . . .252

Stenodactylus . . . .216, 252

Sternaspidae . . 105, 108, 115, 184

Sternaspis . . . . . 108, 184

steudneri, Tropiocolotes .... 252

Sthenelais . . . 106, 113, 116, 128

Sthenelanella . . 106, 116, 128-129

sthenodactylus, Stenodactylus sthenodac-
tylus ...... 252

sthenodactylus sthenodactylus, Stenodac-
tylus ...... 252

Sthenolepis ..... 106, 129

Stichopus . . . . . 117, 1 20

stigmatica, Platysphinx . . . .34

Stolonothrissa .... 329, 337

stroemi, Terebellides . . 108, 116, 194
stylolepis, Lepidasthenia 106, in, 113, 117, 126
subaenea, Glycera ..... 205

subnigra, Coluber rhodorhachis . . 244
Sudani, Taphozous perforatus . 31, 32, 33

swinnyi, Rhinolophus .... 36

swirae, Taphozous perforatus . . 32, 33

Syllidae . . . 104, 105, 106, 139

Syllinae ..... 141-144

Syllis . . . 106, in, 112, 141-142, 143
Synelmis . . . 106, 112, 114, 137

syriacus, Ptyodactylus lobatus . .222

Tadarida

tanganicae, Stolonothrissa

tanganyikae, Lironeca

Taphozous

Tarachodes .

46-48, PI. 3
329

329, 335- 337

31-34. 4

374

Tarbophis
Telepsavus .
Tentacularia

tentaculata, Eunice . '
tenuis, Prionospio .
tephrus, Hipposideros caffer
Terebella
Terebellidae .
Terebellides .
Terebellinae .
terrareginae, Lepidasthenia
tessellata, Coronella
tessellata, Natrix .
tessellata, Natrix tessellata
tessellata tessellata, Natrix
tetelensis, Prionospio
Tetrabothrius
tetraura, Lumbrinereis
Tetreres

INDEX

253
. 178

54- 55

107, 112, 161

173
38, 39

108, 112, 198
105, 108, 194-198

108, 116, 194
195-198
. 125

243

243, 249, 253
243, 249, 253

105, 107, 116, 171-173
54. 62

343
192

Thalassia . .

Thalenessa

Tharyx

thebaica, Nycteris .

125, 128, 156, 168, 178, 187

106, 129

108, 180

33. 34

thebaica, Nycteris thebaica ... 34
thebaica labiata, Nycteris . 34-35

thebaica thebaica, Nycteris . . 34

Thelenota . . . 117, 120

Thelepinae . IQ5

Thormora . . . 106, 112, 113, 126

tibiana, Eunice ..... 34 2
tread welli, Prionospio . . 173

Trewavas, E. . 359-36i

Triaenops . . 4 1 . P'- 3

Trichobranchinae . . . 194

tricolor, Chrysopteron .... 44
tricolor, Myotis . . . 43. 44. 45
tricolor, Rutilus . . 360

tricolor, Selysius . . -44

tridens, Asellia tridens . 40, 41

tridens tridens, Asellia . 40, 41

tridentatus, Lyreidus . 3. 4. 7. 9, 13

trilobata, Amaea . . . 195

trilobata, Amaeana . . . 108, 195

trinchesii, Mastobranchus 108, in, 113, 186
trioculatum, Megalomma 108, in, 116, 200

tristrami, Acanthodactylus . . . 236
tristrami, Acanthodactylus tristrami

236, 250, 252, PI. 4

tristrami, Zootoca ..... 236
tristrami iracensis, Acanthodactylus . 236

tristrami orientalis, Acanthodactylus . 236
tristrami tristrami, Acanthodactylus

236, 250, 252, PI. 4

Trochochaetidae
tropicus, Poecilochaetus
Tropiocolotes
Trypanedenta
Trypanosyllis
tubifex, Eunice
turcica, Lacerta

105, 107, 175-176
107, in, 113, 176

252
106, 142

106, in, 112, 142

107, 140, 161, 343

219

turcicus, Hemidactylus . . 219, 220, 249
turcicus, Hemidactylus turcicus 219-22O, 252
turcicus turcicus, Hemidactylus 219-22O, 252
turcius, Hemidactylus . . .219

typha, Pista . . . 108, 113, 197, 198

typha, Terebella . .198

Typosyllis . . . 106, 107, in, 112,

113, 114, 142-144
tytius, Dynastes ..... 92

ubianensis, Lysilla
uncinata, Dispio
uncinata, Hydroides
unibranchia, Pista
unicornis, Nematonereis
unifasciata, Neanthes
unifasciata, Nereis
uniformis, Exogone
uschakovi, Mercierella

108, 113, 116, 195

. 167

108, 202

197
107, 112, 113, 162

. 107, 148

107, 148

106, in, 140

203

"3. "7.
108,

validobranchiata, Eunice

Varanidae

Varanus . . . .

variabilis, Conchoecia

variegata, Glauconycteris variegata

variegata, Pseudonereis . . 107,

variegata phalaena, Glauconycteris

variegata variegata, Glauconycteris

variegatus, Lithobius

variopedatus, Chaetopterus

vastus, Siphonosoma

vermicularis, Serpula

Vermiliopsis .

vertagus, Cerithium

verugera, Exogone

vesiculosum, Branchiomma

vesiculosum, Megalomma

vesuvianus, Lithobius

viaderi, Notosceles

vinogradovi, Pista

Viperidae j

viridis, Eulalia ....

vitjazi, Conchoecia 259, 269, 275-278,

vitrarius, Spiochaetopterus

vitrarius, Telepsavus

vittata, Mabuia ....

vittata, Mabuya . . . 243,

vittatus, Scincus .

vosseleri, Lithobius

vulgaris recticrista, Chamaelo .

343

253

253
259, 264

42-43
112, 151

42
42-43

302

107, 177
125, 126

1 08, 203

112, 204

117, 160
106, 140

200

108, 113, 200-201

197

253
106, 134
281, 282

. 178

243
249, 253

243

304

232

walhallae, Dynastaspis . . 90-92, 97

watsonana, Eremias guttulata . .252

weberi, Paralacydonia . 107, 116, 155-156

weddelli, Leptonychotes . 53, 54, 55, 64

Werner, Y. L. . . 213-256, Pis. 1-6

wireni, PSosane . . . 108, 116, 194

wrens, Chestnut-shouldered . . 313-328

INDEX 375

Zamenis ...... 244 zeregi, Pseudophoxinus .... 359

Zelandicus, Lithobius .... 303 zeylanica, Psammolyce . 106, in, 113, 128

zeregi, Leuciscus ..... 360 zeylanica, Sthenelais . 106, 113, 116, 128

zeregi, Paraphoxinus .... 360 Zootoca ...... 236

zeregi, Phoxinellus . . . 359, 360